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The  Lichens  of 
Long  Island,  New  York: 

A  Vegetational 
and  Floristic 
Analysis 

Irwin  M.  Brodo 

Holder,  Graduate  Student  Honorarium  for  1959  and  1960 
New  York  State  Museum  and  Science  Service 


I 


BULLETIN  NUMBER  410 
NEW  YORK  STATE  MUSEUM  AND 
SCIENCE  SERVICE 


The  University 
of  the  State 
of  New  York 

ALBANY,  NEW  YORK  — - - 

The  State 
Education 
Department 


FEBRUARY  1968 


The  Lichens  of 
Long  Island,  New  York: 

A  Vegetational 
and  Floristic 
Analysis 

Irwin  M.  Brodo 

Holder,  Graduate  Student  Honorarium  for  1959  and  1960 
New  York  State  Museum  and  Science  Service 


ALBANY,  NEW  YORK 


The  University 
of  the  State 
of  New  York 


The  State 
Education 
Department 


FEBRUARY  1968 


Digitized  by  the  Internet  Archive 
in  2017  with  funding  from 
IMLS  LG-70-15-0138-15 


https://archive.org/details/lichensoflongisl4101brod 


The  Lichens  of  Long  Island,  New  York: 

A  Vegetational  and  Floristic  Analysis1 

by  Irwin  M.  Brodo2 
ABSTRACT 

the  lichen  vegetation  of  long  island  is  discussed  in  broad 
perspective,  yet  with  considerable  detail,  in  an  attempt  to  present  a 
relatively  complete  picture  of  an  important  segment  of  the  North  Ameri¬ 
can  east  coast  lichen  flora.  A  tloristic  list  based  on  complete  collections 
made  throughout  Long  Island  and  some  adjacent  islands  is  supplemented 
by  a  number  of  investigations  of  local  problems  in  lichen  ecology. 

The  ecological  studies  consist  of  transect  analyses  along  the  island’s 
north  shore,  transplant  experiments  concerning  the  vertical  distribution 
of  corticolous  species  as  well  as  the  city  effect,  analyses  of  the  present 
distributions  of  various  species  by  vegetation  type,  and  observations  on 
succession  and  related  phenomena  in  terricolous,  saxicolous,  and  cortico¬ 
lous  communities. 

A  habitat  classification  is  used  to  group  assemblages  of  lichens  into 
“communities.”  Some  discussion  is  presented  on  the  relative  merits  of  such 
a  loose  classification  as  compared  with  a  more  formal  lichen  “union”  or 
"association”  system  used  by  many  European  workers. 

A  consideration  of  some  of  the  environmental  factors  influencing 
lichen  microdistributions  is  presented  along  with  some  supporting  meas¬ 
urements  and  correlations,  but  no  extensive  work  along  these  lines  is 
pursued. 

The  effect  of  New  York  City  on  Long  Island  lichen  distributions  is 
discussed.  Empirical  data  and  theoretical  considerations  are  used  in  con¬ 
cluding  that  the  lichen  distributions  are  influenced  by  air  pollution  as 
well  as  city-induced  drought,  with  the  former  acting  over  longer  distances 
than  the  latter. 

Placing  the  Long  Island  lichen  flora  into  phytogeographic  perspec¬ 
tive  involved  setting  up  a  scheme  of  “elements”  and  “subelements”  for 
eastern  North  America  into  which  the  lichens  could  be  fit.  The  presence 
of  Long  Island  species  in  Asia  and  Europe  was  noted  and  consideration 
was  given  to  problems  of  migration  and  vicariism. 

The  lichen  flora  consists  of  261  species.  Keys  to  the  identification 
of  these  species,  including  keys  to  sterile  material,  precedes  an  extensive 
annotated  list.  Included  under  each  species  in  this  list  is  reference  to 
material  seen,  notes  on  habitat  ecology,  a  statement  on  North  American 
and  world-wide  distribution,  and  where  necessary,  notes  on  nomenclature, 
morphological  and  chemical  variation,  and  closely  related  and/or  con¬ 
fusing  species. 

'Manuscript  submitted  for  publication  January  4,  1966. 

2Curator  of  Lichens,  National  Museum  of  Canada,  Ottawa.  Ontario. 

iii 


Three  species  are  described  as  new:  Polyblastiopsis  quercicola, 
Pertusaria  subpertusa,  and  Lepraria  zonata.  In  addition,  three  new  com¬ 
binations  are  made:  Micarea  prasina  var.  sordidescens  (Nyl.)  Brodo, 
Parmelia  michauxiana  var.  laciniata  (Hale)  Brodo,  and  Buellia  curtisii 
(Tuck.)  Imsh.  in  Brodo. 


iv 


Acknowledgements 

O 

This  work  was  begun  in  1959  with  the  help  of  an  honorarium  from 
the  New  York  State  Museum  and  Science  Service  and  was  continued 
through  1960  under  the  same  auspices.  In  1961-62  the  work  was  con¬ 
tinued  with  the  aid  of  a  graduate  fellowship  from  the  National  Science 
Foundation,  which  provided  the  opportunity  for  uninterrupted  research 
to  complete  the  study.  The  present  monograph  is  based  on  a  dissertation 
in  partial  fulfillment  of  the  requirements  for  a  Ph.D.  degree  from  the 
Department  of  Botany  and  Plant  Pathology  at  Michigan  State  University, 
East  Lansing. 

The  encouragement,  help,  and  guidance  of  Dr.  Eugene  Ogden  during 
this  period  is  especially  appreciated. 

The  skilled  and  dedicated  help  of  Dr.  Henry  Imshaug  was  my  great¬ 
est  asset  throughout  my  studies.  His  guidance,  advice,  and  good  humor 
were  truly  an  inspiration.  Dr.  John  Cantlon's  many  critical  comments  and 
valuable  suggestions  are  greatly  appreciated.  Drs.  Ervin  Barnes,  Edward 
Cantino,  Roland  Fischer,  and  the  late  Dr.  Philip  Clark,  all  contributed 
suggestions  and  comments  on  the  manuscript.  Dr.  Mason  E.  Hale,  Jr.,  in 
the  final  reading  of  the  manuscript,  offered  many  valuable  comments  and 
unpublished  data  for  which  I  am  very  grateful. 

I  especially  would  like  to  thank  Mr.  Roy  Latham,  not  only  for  lend¬ 
ing  me  his  entire  lichen  collection  which  was  of  such  fundamental 
importance  to  this  work,  but  for  his  amiable  and  informative  letters 
concerning  the  Long  Island  of  past  years,  and  for  his  companionship  on 
several  exciting  and  fruitful  field  trips.  Of  the  many  Long  Island  residents 
and  naturalists  who  led  me  to  rich  or  relatively  inaccessible  collecting 
areas,  I  would  like  to  extend  special  thanks  to  Mr.  Leroy  Wilcox,  Mr. 
Gilbert  Raynor,  and  Miss  Linda  Quinby.  Dr.  George  Woodwell's  help  in 
securing  a  room  and  laboratory  facilities  at  Brookhaven  National  Labora¬ 
tory  is  greatly  appreciated,  as  is  Mr.  Frank  McKeaver’s  hospitality  and 
guidance  on  Nantucket  Island. 

My  most  sincere  gratitude  goes  to  my  very  patient  and  tireless  wife, 
Fenja,  for  her  many  and  varied  assistances. 

Thanks  are  due  to  the  following  lichenologists  for  identifying  or 
verifying  Long  Island  material  in  their  special  fields  of  interest;  T.  Ahti 
(Cladina),  W.  L.  Culberson  (Cetraria),  F.  Erbisch  (Chaenotheca),  A.  W. 
Evans  (Cladonia),  M.  E.  Hale  (Parmelia),  W.  Harris  (Polyblastiopsis, 
Leptorhaphis),  A.  Henssen  (Placynthium),  A.  W.  C.  T.  Herre  (Usnea), 
G.  Howard  (Ochrolechia),  I.  M.  Lamb  (Stereocaulon),  A.  H.  Magnusson 
(Ramalina),  E.  D.  Rudolph  (Caloplaca),  H.  Sierk  (Leptogium),  D.  Swin- 
scow  (Porina),  W.  Weber  (Acarospora),  C.  M.  Wetmore  (Nephroma).  Mr. 
W.  D.  Margadant  of  the  Hunt  Library  kindly  helped  me  with  the 
Latin  diagnoses. 


v 


Contents 


PAGE 

INTRODUCTION . 1 

General . 1 

History . 1 

DESCRIPTION  OF  LONG  ISLAND . 5 

Geography . . 5 

Geology . 5 

Climate . 6 

Vegetation  Types . 8 

HABITAT  ECOLOGY . 17 

General  Methods . 17 

1.  Collection  data . 17 

2.  Statistical  studies . 17 

3.  Transplant  experiments . 19 

Substrate . 20 

1.  Texture . 22 

2.  Moisture-holding  capacities . 22 

3.  Stability . 28 

4.  Chemical  composition . 29 

Climate . 31 

1.  Illumination  and  temperature . 31 

2.  Moisture . 32 

Vertical  Distribution . 34 

Succession . 37 

Species  Composition  within  Habitats . 45 

CITY  EFFECT . 63 

FLORISTIC  ELEMENTS . 65 

Introduction . 65 

The  Classification  of  Elements . 65 

Summary  of  Significant  Features . 77 

Discussion . 79 

Summary . 89 

THE  LICHEN  FLORA . 91 

Collections. . 91 

Additional  Specimens  Examined . 107 

Taxonomy . 107 

1.  Species  concept . 107 

2.  Ecological  forms . 109 

3.  Infraspecific  taxa . Ill 

4.  Keys  and  annotated  list . Ill 

vii 


PAGE 


GENERAL  DISCUSSIONS . 267 

Distribution  of  Lichens  on  Long  Island . 267 

1.  Substrate . 267 

2.  Climate . 268 

3.  Vegetation  type . 269 

SUMMARY  AND  CONCLUSIONS . 279 

Habitat  Ecology  and  Lichen  Communities  ....  279 

Lichen  Distributions . 280 

The  City  Effect . 280 

The  Lichen  Flora . 280 

APPENDIX  A  —  Long  Island  Collectors . 282 

APPENDIX  B  —  Glossary . 284 

I.  Morphological  and  Ecological  Terms  ....  284 

II.  Chemical  Terms . 295 

APPENDIX  C  —  Checklist  of  the  Lichens  of  Long  Island  .  .  300 

LITERATURE  CITED . 305 


Illustrations 

FIGURE  PAGE 

1 .  Soil  types . 4 

2.  Original  vegetation . 4 

3.  Precipitation . 4 

4.  Temperature . 7 

5.  Average  annual  number  of  days  of  dense  fog  ...  7 

6.  Relative  humidity . 7 

7.  Sand  dune  vegetation  on  south  shore  dunes . 10 

8.  Sand  dunes  and  sand  plains  .  .  .  near  Montauk  .  .  10 

9.  A  portion  of  a  sand  plain  community  .  .  .  .10 

10.  North  shore  bluffs . 10 

11.  Pine-oak  forest . 14 

12.  Black  oak  forest . 14 

13.  Gravel  pit  bog . 14 

14.  A  sheltered  gravel  beach;  habitat  of 

Verrucaria  microspora . 14 

15.  Collection  localities . 18 

16.  Population  changes  in  a  corticolous  lichen  community: 

non-directional  shifts . 38 

viii 


FIGURE 

PAGE 

17. 

Population  changes  in  a  corticolous  lichen  community: 

directional  shifts . 

39 

18. 

East-west  transect  and  transplant  localities 

41 

19. 

Bark-borer . 

46 

20. 

Arctic-Boreal  element;  Boreal-Temperate  subelement  . 

68 

21. 

Temperate  element;  North  Temperate  subelement  . 

69 

22. 

Temperate  element;  Appalachian  subelement; 

Appalachian  unit . 

70 

23. 

Temperate  element;  Appalachian  subelement; 

Appalachian-Ozark  unit . 

71 

24. 

Temperate  element;  Appalachian  subelement; 

Appalachian-Great  Lakes  unit . 

72 

25. 

Temperate  element;  Appalachian  subelement; 

Appalachian-Great  Lakes-Rocky  Mountain  unit  . 

73 

26. 

Temperate  element;  Coastal  Plain  subelement  . 

74 

27. 

Temperate  element;  East  Temperate  subelement  . 

75 

28. 

Temperate  element;  Oceanic  subelement  .  .  .  . 

76 

29. 

Phytogeographic  affinities  of  the  Long  Island  lichen  flora 

78 

30. 

Historic  relationships  between  floristic  elements, 

subelements,  and  units  in  eastern  America 

85 

31. 

Long  Island  localities  of  oceanic  species  .... 

93 

32. 

Bog  and  swamp  localities . 

93 

33-41. 

Lichens  found  mainly  in  bogs  and  swamps 

93 

42-52. 

Lichens  found  mainly  in  pine-oak  forests  .... 

95 

53-63. 

Lichens  found  mainly  in  morainal  areas  .... 

98 

64-70. 

Lichens  found  mainly  in  the  humid  “fog  belt’’  region  . 

101 

71. 

An  avoidance  of  the  red  oak  forest . 

103 

72-73. 

The  scattered  distribution  of  two  terrestrial  lichens . 

103 

74-76. 

Lichens  found  mainly  on  sand  dunes  and  sand  plains  . 

103 

77-81. 

Lichens  having  a  maritime  distribution  .... 

104 

82. 

Polyblastiopsis  quercicola  (holotype):  Habit  . 

148 

83. 

Polyblastiopsis  quercicolci  (holotype):  Microscopic 

characters . 

149 

84. 

Pertusciria  sub  pert  usa  (holotype) :  Habit  .  .  .  . 

208 

85. 

Pertusaria  subpertusa  (holotype) :  Spores  and  ascus 

209 

86. 

Lepraria  zonata  (holotype)  .  .  .... 

264 

87. 

Ascocarps  . 

289 

88. 

Thallus  types . 

291 

89. 

Lichen  phycobionts . 

291 

90. 

Some  ascospore  types . 

291 

IX 


Tables 


TABLE 

la.  Bark  characters  of  some  common  Long  Island 

phorophytes . 

2.  Sand  and  soil  pH . 

3.  Vertical  distribution  of  some  corticolous  lichens 

in  red  oak  and  pine-oak  forests . 

4.  Degree  of  similarity  of  the  lichen  vegetation  growing 
on  various  species  of  oak  in  the  red  oak  forest  . 

5.  Coefficients  of  association  of  lichen  vegetation  on  differ¬ 
ent  tree  species  at  base  and  breast  height  quadrats  . 

6.  Phytogeographic  categories  represented  in  the 

Long  Island  lichen  flora . 

7.  Phytogeographic  affinities  of  Long  Island  lichens  . 

8.  European-American  vicarious  subgeneric  taxa  in 

the  Long  Island  lichen  flora . 

9.  Distribution  of  some  common  lichens  in  various 

vegetation  types  on  Long  Island . 


PAGE 

24 

26 

35 

51 

52 

80 

84 

87 

272 


x 


Introduction 


GENERAL 

Eastern  North  America  has  received  more  lichenological  study  than 
any  other  part  of  the  continent.  Such  famous  and  productive  workers  as 
Edward  Tuckerman,  Henry  Willey,  Lincoln  W.  Riddle,  R.  Heber  Howe, 
George  K.  Merrill,  Charles  A.  Robbins,  Alexander  W.  Evans,  and  Guy 
G.  Nearing  devoted  much  of  their  lives  to  the  study  of  northeastern 
lichens.  Yet  with  this  exceptionally  fine  background  of  basic  taxonomic 
knowledge,  no  recent  workers  studied  this  area  using  modern  methods  of 
floristic  analysis  and  taxonomy  until  Gunnar  Degelius  visited  the  United 
States  in  1939  and  published  two  excellent  papers,  one  dealing  with  the 
lichens  of  Maine  (Degelius,  1940)  and  the  other  with  the  lichens  of 
the  Smoky  Mountains  of  Tennessee  (Degelius,  1941).  In  1950,  Hale 
wrote  an  account  of  the  lichens  of  Aton  Forest  in  northeastern  Con¬ 
necticut,  and,  in  1954,  I.  Mackenzie  Lamb  published  a  study  of  the 
lichens  of  Cape  Breton  Island,  Nova  Scotia.  Both  papers  significantly 
added  to  our  knowledge  of  the  northeastern  coast  lichen  vegetation. 
Culberson  (1958a)  reported  on  some  lichens  of  North  Carolina  but  dealt 
only  with  the  pine-inhabiting  vegetation. 

This  paper,  then,  is  mainly  designed  to  contribute  to  our  knowledge 
of  the  eastern  coastal  plain  vegetation,  and,  hy  so  doing,  to  provide  a  link 
between  the  studies  of  the  northern  coastal  regions  and  the  Appalachians. 

The  principles  which  guided  the  research  summarized  here  were 
that  a  vegetation  cannot  be  adequately  written  without  a  thorough 
knowledge  of  the  flora,  and  that  a  flora  cannot  be  understood  without  a 
study  of  the  ecological  and  phytogeographic  factors  which  shaped  it. 
In  a  study  of  this  scope,  it  is  impossible  to  answer  all  or  even  most  of 
the  questions  asked  concerning  relationships  and  factors  involved  in  the 
vegetational  picture.  It  is  my  earnest  hope  that  this  study  will  point  to 
the  many  taxonomic,  ecologic,  and  phytogeographic  problems  still  in 
need  of  clarification  and  solution,  and  will  provide  a  stimulus  for  other 
workers  to  add  to  our  knowledge  in  these  and  related  fields. 

HISTORY 

Long  Island  lichenology  had  its  beginnings  quite  early  in  the  history 
of  American  botany.  Halsey  (1823)  published  a  list  of  lichens  collected 
“in  the  vicinity  of  New  York.”  hut  he  did  not  state  explicitly  that  he 
collected  east  of  the  East  River,  and  there  is  some  doubt  as  to  whether 
he  listed  any  Long  Island  specimens.  Specimens  which  were  collected  in 
Brooklyn  and  Queens  by  George  B.  Brainerd  and  George  D.  Hulst 
during  the  1860’s  may  very  well  be  the  earliest  from  Long  Island.  Their 
collections,  deposited  in  the  Brooklyn  Botanic  Garden  Herbarium,  pro¬ 
vide  a  good  basis  for  reconstructing  the  probable  state  of  the  lichen  vege¬ 
tation  of  eastern  New  York  City  prior  to  urbanization  ( p.  275). 


1 


2 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Among  the  other  collectors  of  Long  Island  lichens  during  the  late  19th 
century  is  Charles  H.  Peck,  who  collected  all  forms  of  plant  life  through¬ 
out  New  York  State  during  his  tenure  as  New  York  State  Botanist.  His 
collections  are  in  the  New  York  State  Museum. 

In  1899,  S.  E.  Jelliffe  published  "The  Flora  of  Long  Island,”  which 
listed  54  lichen  taxa  from  various  parts  of  the  island.  G.  S.  Wood  (1905) 
published  additions  to  the  lichen  flora,  adding  18  taxa  to  Jelliffe’s  list. 
In  1914,  Wood  published  a  list  of  lichens  growing  in  the  vicinity  of 
New  York  City  which  included  many  species  from  Long  Island. 

The  Cold  Spring  Harbor  area  was  fairly  well  botanized,  not  only 
by  Jelliffe  and  Wood,  but  also  by  A.  J.  Grout  in  1900  and  Stanley  A.  Cain 
in  the  1930’s  in  connection  with  the  Long  Island  Biological  Institute  at 
Cold  Spring  Harbor.  Since  that  time,  however,  no  botanical  field  work 
has  been  done  there.  Some  lichens  collected  by  Stanley  Cain  as  part 
of  the  “Flora  of  Cold  Spring  Harbor”  are  represented  in  the  New  York 
Botanical  Garden  Herbarium,  but  no  specimens  collected  by  Jelliffe 
or  Wood  were  seen.  Unfortunately,  the  complete  collection  of  the  Cold 
Spring  Harbor  flora  which  existed  at  one  time  (Cain,  pers.  comm.)  could 
be  located  neither  at  the  Biological  Laboratories  at  Cold  Spring  Harbor 
itself  nor  elsewhere. 

Roy  Latham,  one  of  the  most  versatile,  thorough,  and  knowledgeable 
of  the  Long  Island  naturalists,  began  collecting  lichens  in  1908.  He 
confined  his  collecting  to  eastern  Long  Island,  especially  around  Orient 
Point,  and  rarely  went  as  far  west  as  Manorville.  Latham’s  first  con¬ 
centrated  effort  was  connected  with  his  publication  of  the  “Flora  of 
the  Town  of  Southold,  Long  Island  .  .  .”  in  collaboration  with  S.  H. 
Burnham  (Burnham  and  Latham,  1914-25).  The  Farlow  Herbarium  in¬ 
cludes  many  of  these  old  Latham  specimens  which  had  been  sent  to 
Riddle,  Hasse,  or  Merrill  for  identification.  Since  the  early  1900’s  Latham 
has  collected  about  2000  lichen  specimens,  including  many  rare  species. 
His  is  by  far  the  most  complete  collection  of  lichens  made  on  Long 
Island  previous  to  these  studies.  Mr.  Latham  kindly  provided  his  entire 
collection  for  my  use.  Approximately  %  of  the  collection  are  species 
of  Cladonia. 

The  Cladonia  specimens  were  almost  all  determined  in  duplicate  by 
Alexander  Evans,  with  whom  Latham  carried  on  an  active  correspond¬ 
ence  until  Dr.  Evans'  death  in  1960.  Many  of  Latham’s  collections 
represent  the  only  specimens  collected  of  some  species  rare  on  the  island 
(p.  274).  Mr.  Latham  continues  to  be  active,  and  I  have  had  the  good 
fortune  to  accompany  him  on  several  collecting  trips  in  eastern  Long 
Eland. 

Raymond  Torrey  had  a  strong  interest  in  lichens,  especially  of  the 
New  York  City  area,  and  made  many  collecting  trips  to  Long  Island 
particularly  to  study  the  Cladoniae.  His  interests  were  not  confined  to 
the  genus  Cladonia ,  however,  as  is  evidenced  by  his  paper  on  Long 
Island  rock  tripes  (Torrey,  1933).  The  New  York  Botanical  Garden 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


3 


Herbarium  contains  Torrey’s  Cladonia  collections.  These  specimens  were 
all  identified  by  Evans  and  prepared  for  the  herbarium  by  John  W. 
Thomson  (Thomson,  personal  communication).  It  is  surprising  that  no 
other  genus  of  lichens  is  represented  in  the  Torrey  collections. 

Although  Babette  Brown  Coleman  collected  and  published  on  some 
lichens  from  Montauk  Point  (Brown,  1948),  no  extensive  collecting  other 
than  Latham’s  has  been  made  in  recent  years. 

(A  complete  list  of  Long  Island  collectors  is  presented  in  Ap¬ 
pendix  A.) 


4 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  1.  Soil  types  (after  Cline,  1957).  (a)  excessively  drained  hilly 
soil  ( Plymouth-Haven  Association),  (b)  excessively  drained  sandy 
soil  ( Colton-Adams  Association),  (c)  Bridgehampton  fine  sandy 
loam,  (d)  well  drained,  prairie-type  soil  (Hempstead-Bridgehampton 
Association ) . 

Figure  2.  Original  vegetation,  (a)  red  oak  forest,  (b)  pine-oak  forest, 
and  pine  barrens,  (c)  scarlet-black  oak  forest,  (d)  Hempstead  Plains 
grassland,  (e)  downs  grassland  and  dune  heath. 

Figure  3.  Precipitation,  (a)  Mean  precipitation  for  growing  season. 
May  1  to  Sept.  30;  (b)  Mean  annual  precipitation. 


Description  of  Long  Island 

GEOGRAPHY 

Long  Island  makes  up  the  eastern  extension  of  the  southern  tip  of 
New  York  State,  lying  just  to  the  south  of  the  Connecticut  coast  and 
separated  from  the  mainland  to  the  north  by  Long  Island  Sound  and  to 
the  west  by  the  East  River  and  Manhattan  Island.  Long  Island  is  116 
miles  long  and,  at  its  broadest  point,  is  20  miles  wide.  There  are  several 
smaller  islands  just  off  the  shores  of  Long  Island,  and  these  were  visited 
and  included  in  the  study  wherever  possible.  Included  were  Long  Beach, 
lones  Beach,  Fire  Island,  Westhampton  Beach,  Shelter  Island,  Gardi¬ 
ner’s  Island,  and  Fisher's  Island;  not  included  were  Robins  Island  (a 
small  island  in  Peconic  Bay)  or  Plum  Island,  which  is  quarantined. 

Long  Island,  the  geographical  unit,  is  subdivided  into  four  political 
units:  Kings,  Queens,  Nassau,  and  Suffolk  Counties.  Kings  County  (more 
widely  known  by  its  borough  name  —  Brooklyn)  and  Queens  County 
are  part  of  New  York  City.  Brooklyn  is  very  populous  and,  except  for 
one  or  two  large  parks  and  some  swampy  areas  to  the  south,  is  covered 
to  a  large  extent  with  brick,  concrete,  and  asphalt.  Queens  is  not  quite 
so  built  up  and  still  has  many  areas  of  more  or  less  natural  woods  and 
swamps.  Forest  Park,  in  the  center  of  one  of  the  most  populated  parts 
of  Queens,  and  Alley  Pond  Park,  farther  east,  still  show  the  magnificent 
red  and  black  oaks  (Quercus  rubra 1  and  Q.  velutina )  and  tulip  trees 
(Liriodendron  tulipifera )  which  characterized  the  forests  of  that  area 
prior  to  urbanization. 

Nassau  County  is  a  classical  example  of  suburbia.  Extensive  housing 
developments  occupy  its  central  portion  and  large  estates  are  common  on 
the  north  shore.  Much  of  the  area  is  still  relatively  undisturbed,  especi¬ 
ally  on  the  larger  tracts  of  privately  owned  property  to  the  north. 

The  largest  county  in  size  and  the  smallest  in  population  is  Suffolk 
County.  Although  suburban  developments  are  frequent  along  its  western 
edge,  the  greater  part  of  the  area  is  made  up  of  farmland  and  undeveloped 
pine  barrens.  Potatoes  and  cabbage  are  the  chief  crops  produced.  Resorts 
are  common  along  the  entire  south  shore. 

GEOLOGY 

Prior  to  the  Wisconsin  glaciation  the  entire  area  which  is  now  Long 
Island,  except  for  the  western  corner,  was  under  water  and  was  covered 
by  a  number  of  marine  sediments  (MacClintock  and  Richards,  1936). 
Early  Wisconsin  glaciation  (The  Iowan-Tazewell  complex)  laid  down 
two  morainal  ridges  over  this  sediment.  The  first,  the  Ronkonkoma 
moraine,  which  runs  through  the  center  of  the  island  eastward  to  Mon- 
tauk  Point  and  then  off  the  coast  to  Martha’s  Vineyard  and  Nantucket 

'All  phanerogamic  nomenclature  follows  Fernald  (1950)  unless  otherwise  noted. 

5 


6 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Island,  probably  resulted  from  the  Farmdale  advance  (Flint,  1953).  The 
second,  caused  by  a  readvance  of  the  ice  (the  Iowan  advance)  after  a 
slight  withdrawal,  formed  the  Harbor  Hill  moraine  which  extends  east¬ 
ward  to  Orient  Point,  then  to  Fisher’s  Island,  and  finally  to  Cape  Cod. 
A  third  advance,  the  Tazewell,  overrode  the  Harbor  Hill  moraine  (Flint, 
1953)  and  produced  many  of  the  major  topographic  features  we  now 
see  on  the  north  shore,  such  as  the  bluffs  (figure  10),  bays,  and  inlets 
(Nichols,  1958). 

A  broad  outwash  plain  is  associated  with  each  moraine.  It  is 
especially  extensive  south  of  the  Ronkonkoma  moraine,  where  it  forms 
a  low,  flat,  sandy  plain  southward  to  the  ocean.  Wave  action  and  ocean 
currents  formed  the  off-shore  barrier  beaches,  Fire  Island  being  the 
longest. 

Bedrock  can  be  found  only  at  the  western  edge  of  Long  Island  in 
Astoria  (Queens). 

The  topography  of  Long  Island  is  entirely  glacial  in  origin.  With 
the  exception  of  the  moraines  mentioned  above,  the  land  is  extremely  flat. 
The  highest  point  on  the  island  is  428  feet  above  sea  level  at  High  Hill, 
near  South  Huntington.  Kettle  holes  with  associated  bogs  or  lakes  are 
scattered  throughout  the  island  (Fuller,  1914;  Nichols,  1958). 

The  soils  are  formed  on  glacial  parent  material,  and  are  more  or 
less  sandy,  very  well  drained,  and  usually  fairly  acid  (figure  1).  The 
morainal  areas  are  characterized  by  medium  to  moderately  coarse  tex¬ 
tured  glacial  till  ( Plymouth-Haven  association)  often  bearing  large 
glacial  erratics.  Acid  sandy-loams  with  fairly  good  moisture  capacities 
( Bridgehampton  associations)  lie  to  the  south  of  the  moraines  in  most 
places.  Very  well-drained  and  very  acid,  coarse-textured  gravel  and  sand 
of  the  glacial  outwash  (Colton  and  Adams  associations)  make  up  a  large 
part  of  the  southern  edge  of  the  island.  In  central  Nassau  County  the 
soil  morphology  is  much  like  that  of  a  typical  prairie  (Hempstead- 
Bridgehampton  association).  The  soil  is  well  drained,  highly  acid,  and 
with  a  dark-colored  surface  layer  (Cline,  1957). 

CLIMATE 

The  precipitation  over  the  greater  part  of  the  island  is  approximately 
40  to  50  inches  per  year,  or  about  4  inches  per  month,  except  for  the  dry 
months  of  June  and  July  (figure  3).  Droughts  are  not  uncommon  in 
central  Long  Island.  More  than  once  a  year,  on  the  average,  there  is  a 
“dry  spell”  (a  period  of  at  least  15  consecutive  days,  none  of  which 
receives  0.05  of  an  inch  or  more  of  precipitation).  Approximately  once 
every  2  years  there  is  an  “absolute  drought”  (15  consecutive  days,  none 
of  which  receives  0.01  of  an  inch  of  rain  or  more).  East  of  Three  Mile 
Harbor,  the  rainfall  averages  30  to  40  inches  per  year.  (Data  and  defi¬ 
nitions  kindly  furnished  by  Brookhaven  National  Laboratory  Meteorology 
Group.) 


Figure  5.  Average  annual  number  of  days  of  dense  fog. 

Figure  6.  Relative  humidity.  — - - - - —  8:00  a.m.,  -  noon, 

- 8:00  p.m.  (a)  Aver,  for  July;  (b)  Aver,  for  Jan. 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Temperatures  on  Long  Island  are  rather  mild,  and  differences  are 
slight  from  one  part  of  the  island  to  another.  On  the  average,  the  winter 
temperatures  are  about  the  same  throughout  the  island,  but  are  milder 
than  farther  inland  due  to  the  oceanic  effect.  Summer  temperatures  grade 
from  warmest  in  the  New  York  City  area  to  coolest  at  the  eastern  half 
of  the  island  (U.S.D.A.,  1941)  (figure  4).  At  Brookhaven  National 
Laboratory,  in  central  Long  Island,  the  average  recorded  temperature 
was  65°  F.  between  October  1  and  September  30,  and  40°  F.  between 
October  1  and  April  30.  Temperatures  in  that  area  rarely  go  below 
10°  F.  or  above  90°  F. 

Winds  are  quite  brisk  all  over  the  island.  In  the  central  portions, 
over  half  the  time  winds  are  between  5.6  and  9  m/sec.  ( 1 1  and  20  miles 
per  hour),  with  winds  over  11  m/sec.  occurring  8  percent  of  the  time. 
Montauk  Point,  on  the  eastern  tip  of  the  island,  is  well  known  for  its  high 
winds.  Prevailing  winds  are  from  the  southwest  during  the  summer  and 
the  northwest  during  the  winter. 

Fog  and  mist  are  common  phenomena  on  the  eastern  tip  of  Long 
Island,  particularly  in  the  Montauk  area  (figure  5).  Depressions  in  the 
downs  and  between  the  dunes  where  fog  can  form  create  local  pockets 
of  extremely  high  humidity  in  the  Montauk  region  ( p.  32). 

Almost  every  autumn.  Long  Island  is  subjected  to  violent  storms 
which  originate  as  hurricanes  in  the  Caribbean  and  sweep  up  the  east 
coast.  Most  of  the  storms  do  only  minor  wind  damage  to  the  plant  com¬ 
munities,  but  occasionally  severe  storms  cause  extremely  high  tides, 
violent  winds,  heavy  salt  spray,  and  driving  rains  which  do  considerable 
damage  along  the  coast  and  even  farther  inland,  particularly  on  the 
eastern  tip  of  the  island.  Roy  Latham  (in  a  letter)  relates  how  the 
hurricanes  of  1938  and  1944  completely  flooded  the  beach  at  Orient 
Point  (Long  Beach)  and  swept  away  a  great  quantity  of  vegetation, 
including  all  but  traces  of  the  lichen  flora.  Tides  rose  12  feet  and  even 
the  corticolous  lichens  were  washed  into  the  ocean. 

Trewartha  (1961),  in  his  modification  of  Kbppen’s  classification  of 
climatic  regions,  placed  the  Long  Island  area  into  his  “Daf”  category  — 
indicating  a  humid,  continental  climate  with  warm  summers. 

In  summary,  the  climate  of  Long  Island  is  characterized  by  periodi¬ 
cally  droughty,  warm  summers  and  rainy,  mild  winters.  In  addition  to 
the  normally  warm  and  droughty  summers  are  the  high  winds  and  ex¬ 
cessively  drained  soils,  greatly  increasing  vegetational  drought.  The 
situation  is  somewhat  alleviated  locally  by  moist  on-shore  winds  and  fogs 
in  the  extreme  eastern  part  of  the  island,  where  the  rainfall  is  the  least 
and  the  winds  are  the  highest. 

VEGETATION  TYPES 

When  one  speaks  of  the  “vegetation  of  Long  Island,”  it  must  be 
understood  that  in  many  areas  there  are  two  vegetations  to  be  discussed 
—  that  of  the  present,  and  that  of  the  presettlement  period.  This  is 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


9 


especially  true  in  the  New  York  City  area  and  adjoining  Nassau  County, 
where  urbanization  virtually  eliminated  once  important  and  conspicuous 
vegetation  types  and  left  only  fragmentary  remnants.  For  example,  Forest 
Park,  on  the  Brooklyn-Queens  boundary,  is  the  only  surviving  remnant 
of  a  forest  described  as  having  been  “heavily  wooded  with  large  timber 
of  an  aspect  similar  to  the  timber  of  the  Connecticut  coasts”  (Svenson, 
1936).  As  late  as  1917,  Harper  reported  some  remnant  forests  in  the 
Queens  area  as  constituting  rich  woods  broken  with  streams  and  mead¬ 
ows.  Some  of  the  larger  trees  Harper  listed  as  being  most  abundant  were 
Quercus  velutina,  Q.  alba,  Hicoria  alba  (Carya  tomentosa),  and  Castanea 
dentata,  with  Quercus  coccinea  being  important  in  the  drier  woods  and 
Liriodendron  tulipifera  being  conspicuous  in  the  rich  woods. 

Another  excellent  example  of  this  massive  vegetational  obliteration 
can  be  seen  in  Nassau  County,  in  the  “Hempstead  Plains”  region.  Orig¬ 
inally,  this  area  was  a  16  mile  long  botanical  oddity  —  a  natural  true 
prairie  on  Long  Island.  The  land  was  not  good  for  farming  because 
of  the  dense,  hard  sod,  but  it  was  used  extensively  for  pasture  (Svenson, 
1936).  Hicks  (1892)  wrote  a  detailed  account  of  the  flora  of  the  Hemp¬ 
stead  Plains.  The  broad,  unforested,  gently  rolling  landscape  provided 
a  perfect  situation  for  mass-produced  housing.  After  the  great  expansion 
in  suburban  living  just  after  World  War  II,  many  housing  developments 
arose  on  the  “plains”  such  as  those  in  Levittown,  Garden  City,  and 
Mineola.  At  this  date,  the  only  remnants  of  this  fascinating  vegetation 
type  are  found  on  fragments  of  the  property  adjoining  some  parts  of  the 
Meadowbrook  Parkway  and  parts  of  Mitchell  Air  Force  Base.  It  will 
later  be  pointed  out  that  the  lichen  flora  occurring  on  these  fragments  is 
amazingly  rich  for  such  a  far  western  position  on  Long  Island. 

The  original  vegetation  of  Suffolk  County,  on  the  other  hand, 
although  fragmentary  and  relegated  to  parks  in  some  areas  to  the  west, 
remains  in  a  more  or  less  recognizable  state  (figure  2).  Conard  (1935) 
presented  a  vegetational  analysis  of  the  vegetation  types  of  central  Long 
Island,  giving  them  phytosociological  binomials.  Among  the  most  con¬ 
spicuous  communities  are  the  well  developed  oak  forests  seen  mostly 
on  the  north  shore  (Harper,  1917;  Cain,  1936),  the  pine  barrens  which 
are  well  developed  in  central  Long  Island  eastward  to  Riverhead  (Harper, 
1908;  Britton,  1880),  and  the  heathlike  “downs”  (as  described  by  Taylor, 
1923)  which  are  very  conspicuous  in  the  Montauk  area.  Also  important 
are  the  communities  characteristic  of  the  sand  dunes  (Brodo,  1961a), 
the  Chamaecy paris  bogs  (Bicknell,  1908;  Harper,  1907;  Nichols,  1907; 
Taylor,  1916),  the  red  maple  swamps  (Cain  and  Penfound,  1938),  and 
the  Hempstead  Plains  (Hicks,  1892;  Harper,  1911,  1912;  Cain,  Nelson, 
and  McLean,  1937). 

More  detailed  breakdowns  of  the  plant  associations  have  been  made 
by  many  authors  (Miller  and  Young,  1874;  Jelliffe,  1899;  Taylor,  1915, 
1922;  Grier,  1925;  Conard,  1935;  Svenson,  1936;  Brodo,  1961a).  The 
names  used  in  the  following  descriptions  are  those  most  widely  accepted 


10 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


and  used  by  the  above  authors  and  other  naturalists  in  the  area.  The 
categories  I  used  in  a  previous  paper  (Brodo  1961a),  although  well 
suited  for  describing  central  Long  Island  stands,  had  to  be  somewhat 
expanded  to  be  of  use  in  depicting  the  vegetation  types  throughout  the 
entire  island. 

1.  Dune  grass  —  Beach  Heather  —  Shrub  Savanna  and  Sand  Plains 
(formed  on  dune  sand;  excluding  pine  barren  glades)  (figures  7, 
8).  Dominant  trees:  Pinus  rigida.  Primus  serotina  (both  sparse 
and  usually  stunted).  Dominant  undergrowth  and  ground  cover: 
Ammophila  breviligulata,  Myrica  pensylvanica.  Primus  maritima, 
Arctostaphylos  uva-ursi,  Hudsonia  tomentosa.  Soil:  quartz  dune 
sand  with  little  or  no  organic  matter.  Light2:  unlimited. 

Most  of  the  barrier  beach  on  the  south  shore  and  a  few  small  areas 
on  the  north  shore  are  composed  of  long,  rolling  dunes,  some  still 
moving.  The  best  developed  dunes  and  their  corresponding  vegetation 
can  be  found  along  the  entire  length  of  Fire  Island,  and  magnificent, 
huge,  moving  dunes  can  be  seen  in  the  Napeague-Promised  Land  area 
and  on  the  western  edge  of  Hither  Hills  State  Park,  facing  Napeague 
Harbor.  Trees  are  very  sparse  and  occur  mainly  in  boggy  depressions 
between  the  dunes.  More  exposed  trees  are  almost  always  dwarfed  into 
a  “krummholz”  form. 

Depressions  and  hollows  between  the  dunes  are  termed  “slacks”  or 
“lows”  by  Salisbury  (1952)  for  those  with  or  without  standing  water, 

'Light:  unlimited  —  almost  entire  area  in  open  sunlight;  excellent  —  at  least 
Vn  the  area  in  open  sunlight,  the  rest  in  moderate  shade;  good  —  less  than  V i 
the  area  in  open  sunlight,  the  rest  in  moderate  shade;  fair  —  no  open  sunlight 
falling  on  ground,  but  some  sunlight  filtering  through  the  trees;  poor  —  tightly 
closed  canopy  with  virtually  no  sunlight  reaching  the  ground. 


Figures  7-10.  Vegetation  types. 

7.  Sand  dune  vegetation  on  south  shore  dunes  near  Quogue,  facing 
the  ocean,  consisting  mainly  of  Ammophila  breviligulata,  Hudsonia 
tomentosa,  and  Myrica  pensylvanica. 

8.  Sand  dunes  and  sand  plains  at  Napeague  near  Montauk.  Ground 
cover  is  mainly  Arctostaphylos  uva-ursi  and  Cladonia  (subgenus 
Cladina)  spp.  A  few  scattered  scrub  oaks  (Quercus  ilicifolia)  and 
pine  ( Pinus  rigida)  can  also  be  seen. 

9.  A  portion  of  a  sand  plain  community  showing  dune  grass 
( Ammophila  breviligulata ),  false  heather  (Hudsonia  tomentosa), 
and  the  light  colored  Cladoniae,  mainly  Cladonia  submitis  and 
C.  boryi. 

10.  North  shore  bluffs  overlooking  Long  Island  Sound  (to  the 
left  of  the  picture).  At  the  summit  of  the  bluffs  can  be  seen  a 
portion  of  the  red  oak  forest.  The  trees  on  the  slope  are  mainly 
Primus  serotina. 


12 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


respectively.  They  have  local  conditions  of  high  moisture  and  cool 
temperatures  due  to  their  receiving  runoff  from  surrounding  dunes  and 
persistent  morning  fogs,  coupled  with  cool  air  drainage  and  protection 
from  drying  wind  action.  Salisbury  (1952)  also  points  out  that  such 
areas  may  he  rich  in  soil  nutrients  (as  compared  with  surrounding 
dunes)  due  to  leaching  and  drainage  into  the  hollows  of  minerals  and 
some  organic  matter. 

Dune  grass  (Ammophila  breviligulata)  is  the  most  vigorous  of  the 
dune  plants  and  is  found  throughout  the  area,  with  shruhs  such  as  Myrica 
pensylvanica,  Primus  maritima,  and  Toxicodendron  radicans  growing 
mainly  on  the  lee  sides  of  dunes.  Bearherry  (Arctostaphylos  uva-ursi)  and 
false  heather  (Hudsonia  tomentosa)  are  often  conspicuous  on  more  ex¬ 
posed  areas  between  the  dunes  ( Brodo,  1961a).  Conard  (1935),  whose 
Ammophiletum  breviligulatae,  Hudsonietwn  tomentosi.  Primus  maritima - 
Myrica  carolinensis  (M.  pensylvanica )  association,  and  Pinus  rigida  scrub 
association  all  fit  into  this  vegetation  type,  noted  the  close  similarity  of 
this  community  to  the  dune  communities  of  Europe.  Martin  (1959) 
describes  this  vegetation  type  in  detail,  as  it  occurs  in  New  Jersey  (see 
especially  his  communities  1-2,  8-11,  15-18,  24-29,  44). 

2.  Pine  barrens  (part  of  continuum  segment  A  in  Brodo,  1961a). 
Dominant  trees:  Pinus  rigida,  Quercus  alba,  Q.  coccinea.  Domi¬ 
nant  undergrowth:  Quercus  ilici folia,  Gaylussacia  baccata,  Vac- 
cinium  angustifolium,  V.  vaccilans,  Pteridium  aquilinum.  Soil: 
dune  sand  or  Colton  and  Adams  sandy  loam.  Light:  good  to 
excellent. 

The  wide  expanses  of  pitch  pine  (Pinus  rigida)  and  scrub  oak 
(Quercus  ilicifolia)  which  are  characteristic  of  most  of  central  Long 
Island  have  existed  for  centuries  virtually  unchanged.  George  Washing¬ 
ton  wrote  in  his  diary  on  April  22,  1790,  a  description  of  the  area  he 
saw  as  he  rode  from  Patchogue  to  Coram  and  Setauket.  He  described  the 
area  as  “too  poor  for  cultivation,  being  low  scrubby  oak,  not  more 
than  two  feet  high,  intermixed  with  small  and  ill  thriven  pines”  (Taylor, 
1922). 

Conard  (1935)  states  that  this  basic  community  extends  from 
Newfoundland  (where  it  is  fragmentary)  south  to  Georgia  and  Texas, 
with  Pinus  taeda  and  P.  palustris  replacing  P.  rigida  as  the  dominant. 
Both  his  Pinetum  rigidae  and  Quercetum  ilici foliae  communities  can 
be  placed  here. 

3.  Pine-oak  forest  (continuum  segments  A  and  B  in  Brodo,  1961a) 
(figure  II).  Dominant  trees:  Quercus  alba,  Q.  coccinea,  Pinus 
rigida.  Dominant  undergrowth:  as  in  pine  barrens,  with  Q.  ilici¬ 
folia  sparse  except  in  glades.  Soil:  Bridgehampton  sandy  loam. 
Light:  good. 

This  vegetation  type  is  little  more  than  an  older,  more  mature  pine 
barren.  The  three  dominant  trees  are  the  same  in  both,  but  the  order  of 
abundance  is  different  in  the  pine-oak  forest  with  the  appearance  of 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


13 


Quercus  velutina.  The  soil  is  better  developed  with  more  organic  matter, 
although  the  ground  vegetation  is  essentially  the  same.  The  trees  are 
generally  older,  taller,  and  straighter.  Sparrow  and  Woodwell  (1962) 
presented  a  good  description  of  this  vegetation  type  in  their  description 
of  a  radiation  study  area  at  Brookhaven  National  Laboratory  in  central 
Long  Island.  The  Quercetum  velutinae  as  described  by  Conrad  (1935) 
belongs  here  and  can  also  be  applied  to  the  scarlet-black  oak  woods 
discussed  below. 

4.  Scarlet-black  oak  forest  (continuum  segment  C  in  Brodo,  1961a) 
(figure  12).  Dominant  trees:  Quercus  coccinea,  Q.  velutina, 
Q.  alba.  Dominant  undergrowth:  as  in  pine-oak  forest.  Soil: 
Bridgehampton  sandy  loam.  Light:  good. 

Again,  we  have  a  slightly  older,  more  mature  forest  of  basically  the 
same  structure  as  the  previous  vegetation  types.  Pinus  rigida  becomes 
relatively  unimportant  here  with  the  increasing  importance  of  Quercus 
velutina. 

5.  Red  oak  forest  (continuum  segment  D  in  Brodo,  1961a).  Dom¬ 
inant  trees:  Quercus  velutina,  Q.  rubra,  and  locally,  Q.  prinus. 
Dominant  undergrowth:  Viburnum  acerifolium,  Smilax  glauca, 
Vaccinium  sp.,  Parthenosissus  quinquefolia.  Soil:  Plymouth- 
Haven  loam,  generally  with  a  good  humus  accumulation,  on 
glacial  till.  Light:  fair  to  poor. 

The  red  oak  forest  extends  all  along  the  north  shore  and  includes 
parts  of  the  Sag  Harbor  region.  It  is  this  vegetation  type  which  originally 
covered  much  of  the  New  York  City  area  and  which  was  described  by 
Harper  (1917).  Ground  cover  in  the  present  stands  is  usually  sparse, 
except  in  some  local  spots  where  Smilax  species  and  Rubus  species  grow 
in  dense  thickets. 

Conard’s  (1935)  Quercetum  kalmietosum  and  Quercetum  prini  both 
seem  to  fit  best  here.  Where  the  soil  is  moist,  Fagus  begins  to  come  and 
replace  the  oaks  (Conard,  1935). 

6.  Beech-oak  forest.  Dominant  trees:  Fagus  grandifolia,  Quercus 
rubra,  Acer  rubrum.  Dominant  undergrowth:  very  sparse.  Soil: 
Plymouth-Haven  loam  with  much  humus  on  till.  Light:  poor. 

A  few  small,  isolated  areas  near  the  eastern  tip  of  Long  Island  bear 
remnants  of  some  of  the  oldest  vegetation  on  the  North  American  east 
coast.  These  forests  of  old  beech  and  oak  trees  can  be  found  on 
Gardiner’s  Island,  near  Montauk  Point,  and  on  Shelter  Island  (Taylor, 
1923). 

7.  Downs.  Dominant  trees:  Primus  serotina,  Amelanchier  inter¬ 
media.  Dominant  undergrowth:  Myrica  pensylvanica.  Primus 
maritima.  Dominant  groundcover:  Andropogon  scoparius.  Soil: 
Colton  and  Adams  sandy  loam.  Light:  unlimited. 

Norman  Taylor  (1923)  wrote  a  detailed  account  of  the  grasslands 
of  the  Montauk  region.  As  far  as  records  show,  the  area  was  always  a 
grassland  devoid  of  any  substantial  forest  cover.  Primus  serotina  is  the 


14 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


only  conspicuous  tree  in  the  entire  grassland  area,  and  it  is  of  very 
scattered  occurrence.  Amelanchier  intermedia  also  occurs  in  a  few 
groves.  Shrubs  are  scattered  throughout  the  area.  Taylor  (1923)  stated 
that  “wind  is  unquestionably  the  most  important  (factor)  in  maintaining 
the  area  as  a  grassland.” 

This  community  is  called  the  Andropogon  scoparii  in  Conard  (1935). 

8.  Hempstead  Plains  grassland.  Dominant  tree:  Primus  serotina. 
Dominant  shrub:  Myrica  pensylvanica.  Dominant  ground  cover: 
Andropogon  scoparius.  Soil:  Hempstead-Bridgehampton  sandy 
loam.  Light:  unlimited. 

A  great  deal  of  work  has  been  done  on  the  vegetation  of  the  Hemp¬ 
stead  Plains  (p.  9).  It  is  considered  by  most  workers  to  be  a  true 
“natural  prairie,”  i.e.,  a  stable  grassland  community.  The  long  stretches 
of  Andropogon  scoparius  are  only  occasionally  broken  by  isolated  black 
cherry  trees  or  bayberry  bushes.  Wind  was  probably  not  an  important 
factor  in  the  development  of  the  prairie  here  as  it  was  with  the  very 
similar  Montauk  downs,  since  Hempstead  Plains,  in  central  Nassau 
County,  is  not  an  especially  windy  area.  Hicks  (1892)  claimed  that 
excessive  drainage  plus  the  thinness  of  the  surface  soil  and  general 
climate  determined  the  character  of  the  flora  of  the  plains. 

The  soil  is  made  up  of  tight,  matted  sod  with  sandy,  eroded  areas 
occurring  wherever  the  sod  had  been  broken.  This  dense  sod,  almost 
too  hard  to  plow  through  and  too  dense  to  allow  tree  roots  to  penetrate, 
probably  prevented  subsequent  forestation  by  local  trees  (Svenson,  1936). 
Conard  (1935)  called  this  community  the  Andropogon  Hempsteadi. 

9.  Bogs.  Dominant  trees:  Chamaecy paris  thyoides,  Acer  rubrum, 
Nyssa  sylvatica.  Dominant  shrubs:  Vaccinium  corymbosum, 
Toxicodendron  vernix  (L.)  Kuntze.  Dominant  ground  cover: 
Sphagnum  spp.,  Vaccinium  macrocarpon,  V.  oxycoccos.  Wood - 
wardia  virginica.  Soil:  wet,  acid  sand  grading  into  acid  peat. 
Light:  excellent  to  poor,  depending  on  canopy  development. 

Figures  11-14.  Vegetation  types. 

11.  Pine-oak  forest  at  Brookhaven  National  Laboratory  in  central 
Long  Island,  dominated  by  Pinus  rigida,  Quercus  alba  and  Q. 
coccinea. 

12.  Black  oak  forest  near  Manorville  in  central  Long  Island,  with 
tall  Quercus  velutina  and  Q.  coccinea  and  an  undergrowth  of 
Vaccinium  spp. 

13.  A  small  gravel  pit  bog  near  the  south  shore  at  Eastport,  sur¬ 
rounded  by  Pinus  rigida.  Lush  stands  of  Cladonia  atlantica  were 
found  here. 

14.  A  sheltered  inlet  and  gravel  beach  on  Shelter  Island  (Ram 
Island  Neck)  which  was  the  habitat  of  a  collection  of  Verrucaria 


microspora. 


16 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


White  cedar  swamps  at  one  time  were  abundant  all  along  the  south 
shore  at  the  heads  of  tidal  streams  and  salt  marshes  (Harper,  1907; 
Nichols,  1907;  Bichnell,  1908;  Taylor,  1916).  Heusser  (1949),  who  pre¬ 
sented  the  history  of  such  an  “estuarine  bog”  from  the  nearby  New 
Jersey  coast,  stated  that  rising  sea  level,  ditching  (with  the  subsequent 
influx  of  brackish  water),  and  fires  caused  the  disappearance  of  the 
cedars  in  that  area.  Similar  conditions  probably  occurred  on  the  Long 
Island  coast.  In  addition,  with  the  spread  of  suburbanization,  almost  all 
the  cedars  in  Nassau  County  were  harvested  and  most  of  the  swamps 
filled  in  to  provide  space  for  the  ever-extending  highways.  Although 
there  are  still  some  fragmentary  estuarine  bogs  in  the  Babylon  area,  the 
best  developed  bogs  are  those  farther  east  and  inland  which  were  formed 
in  glacial  depressions  and  are  surrounded  by  pine  or  pine-oak  forests 
(figure  13).  In  the  Manorville  region,  some  bogs  were  extensively  culti¬ 
vated  for  cranberries,  but  few  are  still  in  use.  The  soil  is  very  acid  and 
provides  a  good  habitat  for  bog  plants  such  as  Vaccinium  macrocarpon, 
V .  oxycoccos,  Drosera  spp..  Lycopodium  spp.,  Sarracenia  purpurea,  and 
Utricularia  spp. 

The  white  cedar  swamps  in  various  stages  of  development  make  up 
the  Chamaecypareum  thyoidis,  Chamaedaphnetum  calycidatae,  and  Vac- 
cinietum  corymbosi  of  Conard  (1935).  An  otherwise  similar  commu¬ 
nity,  but  without  white  cedar,  has  been  called  the  Aceretum  rubri  and  is 
discussed  next. 

10.  Red  maple  swamp.  Dominant  trees:  Acer  rubrum,  Nyssa  syl- 
vatica.  Dominant  shrubs:  Clethra  alnifolia,  Viburnum  dentation, 
Vaccinium  corymbosum.  Ground  cover:  sparse;  Osmunda  sp., 
Sphagnum  spp.  Light:  fair  to  poor. 

In  wet  areas  not  suited  for  white  cedar,  red  maple  swamps  become 
established.  They  are  common  throughout  the  island.  Cain  and  Penfound 
(1938)  described  and  discussed  this  vegetation  type  in  considerable 
detail,  referring  to  it  as  the  Aceretum  rubri  (including  both  the  Aceretum 
rubri  and  the  Aceretum  osmundaceum  of  Conard  [1935]). 

It  can  easily  be  seen  that  vascular  vegetation  and  soil  type  are 
strongly  correlated  (compare  figures  I  and  2).  The  red  oak  forests  are 
largely  confined  to  the  Plymouth-Haven  soils,  the  pine-oak  forests  remain 
closely  correlated  with  the  Bridgehampton  sandy  loam,  and  the  pine 
barrens  are  best  developed  on  the  Colton  and  Adams  coarse  sands.  The 
Hempstead  Plains  grasslands  are  confined  to  the  Hempstead-Bridge- 
hampton  soil  association,  which  is  considered  to  have  been  formed  under 
grassland  vegetation  (Cline,  1957).  The  dune  and  down  vegetation  of 
the  south  shore  occurs  largely  on  windblown  dune  sand. 


Habitat  Ecology 


GENERAL  METHODS 

1.  Collection  data.  Many  important  ecological  notes  on  particular 
species  were  gleaned  from  label  data  of  individual  collections.  For  very 
rare  species,  these  were  often  the  only  data  available  other  than  my  field 
notes.  On  my  own  collections,  substrate  was  noted  as  accurately  as  pos¬ 
sible  for  each  specimen  (e.g.,  the  phorophyte  species  in  the  case  of 
corticolous  lichens).  If  the  phorophyte  species  could  not  be  determined 
in  the  field,  I  collected  a  portion  of  a  branch  or  twig  with  the  lichen  for 
later  identification.  Height  above  ground  was  noted  along  with  other 
parameters,  such  as  exposure  in  relation  to  a  body  of  water,  a  road,  or  a 
farm,  if  thought  to  be  locally  important.  For  each  locality  I  recorded 
the  general  light  conditions,  the  dominant  tree  layer,  shrub  layer,  and 
ground  cover. 

Since  collections  are  not  unbiased,  label  data  are  of  no  use  in 
statistical  studies  and  only  limited  information  can  be  gathered  from 
this  source  concerning  host  specificity,  vertical  distribution,  etc.  Label 
data  are  of  greatest  use  in  determining  where  a  species  can  occur,  i.e.,  the 
substrate  potential;  and  never  where  it  cannot  occur,  i.e.,  the  substrate 
limits.  Label  data  may  indicate  trends  and,  where  the  number  of  collec¬ 
tions  is  large  and  the  ecological  limits  small,  certain  tentative  conclusions 
may  be  drawn. 

2.  Statistical  studies.  There  are  many  ecological  phenomena  which 
can  be  studied  adequately  only  through  the  use  of  unbiased  sampling  and 
statistical  analyses.  Questions  pertaining  to  substrate  specificity,  the 
range  and  frequency  of  species  in  different  wooded  stands,  the  effects 
of  New  York  City  on  lichen  distribution,  and  the  vertical  zonation  of 
corticolous  lichens  were  all  approached  statistically  with  the  following 
methods. 

Lichen  sampling  was  carried  out  in  two  areas,  one  in  central  Long 
Island  in  1959,  and  one  on  the  north  shore  in  1961.  Different  sampling 
methods  were  employed  in  the  two  studies,  but  since  both  involved  un¬ 
biased  samples  of  small  areas,  the  data  should  be  comparable. 

In  the  first  case,  1 1  stands  in  central  Long  Island  were  sampled 
using  a  modification  of  the  “random  pairs”  method  of  Cottam  and 
Curtis  (1949).  The  method  has  been  fully  described  in  previous  papers 
(Brodo,  1961a;  Culberson,  1955a;  Hale,  1955a).  Briefly,  the  method 
consisted  of  selecting  pairs  of  trees  at  prearranged  intervals  along  a 
randomly  selected  transect  line  until  20  pairs  (40  trees)  were  exam¬ 
ined.  On  each  tree,  two  quadrats  were  studied,  one  from  the  ground 
level  to  a  height  of  30  cm,  and  another  40  cm  high,  centered  at  1.3m 
(breast  height).  Each  quadrat  encircled  the  trunk.  The  stands  sampled  in 
the  1959  study  ranged  from  pine  barrens  to  red  oak  forests. 

The  second  sampling  study  was  done  in  1961  in  the  red  oak  forests 
along  the  north  shore.  Twelve  stands  were  sampled  along  an  east-west 


17 


18 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


«  % 


Figure  15.  Collection  localities. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  19 

transect  starting  at  Forest  Park  in  New  York  City  and  going  eastward 
to  Shoreham  in  central  Long  Island  (figure  18). 

For  the  purpose  of  this  study  it  was  desirable  to  limit  the  survey  to 
red  oak  stands  of  fairly  uniform  composition.  Due  to  the  uneven  topog¬ 
raphy  of  the  morainal  north  shore,  vegetation  appeared  very  patchy 
and  areas  of  more  or  less  uniform  tree  composition  were  small.  For 
this  reason  the  random  pairs  method  used  in  the  previous  study  was 
unsuitable,  since  it  covered  too  much  territory  and  the  vegetation  within 
each  sample  would  have  been  too  diverse. 

Instead,  a  spiral  sampling  technique  was  employed.  The  system 
simply  consisted  of  choosing  a  point  in  the  center  of  the  area  to  be 
sampled  and  working  in  an  ever-increasing  spiral,  examining  all  encoun¬ 
tered  trees  until  50  had  been  studied.  This,  then,  is  essentially  a  100 
percent  sample  of  a  very  small  area.  The  selection  of  the  starting  point 
in  each  stand  was  made  to  center  specifically  in  the  greatest  concentration 
of  red  oaks,  regardless  of  the  lichen  population.  This  entirely  non-random 
selection  of  stands  is  valid  since  it  is  not  the  tree  vegetation  which  is 
under  study,  but  rather  the  epiphytic  vegetation  of  those  trees.  By  select¬ 
ing  stands  for  a  certain  tree  composition,  the  important  variable  of  forest 
type  is  largely  eliminated  and  the  epiphytic  vegetation  within  the  stand 
can  still  be  sampled  in  an  unbiased  manner. 

On  each  tree,  two  cylindrical  quadrats,  delimited  exactly  as  in  the 
1959  study,  were  examined.  Neither  dead  trees  nor  any  that  were  less 
than  10  cm  in  diameter  at  breast  height  (dbh)  or  inclined  at  an  angle 
of  more  than  10°  were  considered.  Data  sheets  were  constructed  to 
include  about  25  common  lichens,  all  of  which  could  be  identified  in  the 
field  without  question.  The  species  and  dbh  of  each  tree  as  well  as  the 
presence  in  each  quadrat  of  any  listed  lichen  were  recorded.  Cover  was 
not  noted,  but  the  direction  of  exposure  of  each  species  was  recorded  by 
noting  its  presence  for  each  of  eight  compass  points. 

The  lichens,  as  a  rule,  were  easily  identified  in  the  field  with  a  hand 
lens,  although  occasionally  chemical  tests  were  performed  on  the  thalli 
with  potassium  hydroxide,  p-phenylenediamine,  or  hypochlorite  solution 
for  confirmation.  The  phorophyte  species  were  often  more  difficult  to 
determine,  perhaps  owing  to  the  apparent  wide  occurrence  of  hybridiza¬ 
tion  in  the  area  among  members  of  the  black  oak  group  (Quercus  velu- 
tina,  Q.  rubra,  and  Q.  coccinea).  If  the  tree  under  study  was  judged  to 
be  a  hybrid,  the  two  putative  parent  species  were  listed  in  place  of  a 
single  species  name  (e.g.,  Quercus  rubra  X  coccinea).  Previously  (Brodo, 
1961a),  these  three  members  of  the  black  oak  group  were  considered 
collectively  under  the  name  of  Quercus  velutina.  As  will  be  pointed  out 
later,  the  epiphytic  lichen  populations  on  the  three  species  are  very 
similar. 

3.  Transplant  experiments.  In  an  effort  to  clarify  some  of  the  eco¬ 
logical  factors  governing  lichen  distributions,  some  transplant  experi¬ 
ments  using  corticolous  lichens  were  carried  out.  The  methods  employed 


20 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


were  fully  described  in  a  previous  paper  ( Brodo,  1961b)  but  a  brief 
account  will  be  presented  here,  incuding  a  few  modifications  and  im¬ 
provements  which  were  used  in  the  latest  experiments. 

Using  a  steel  punch  (hereafter  referred  to  as  a  “bark-borer”)  con¬ 
sisting  of  a  hole-saw  blade  bevelled  on  the  outside  to  a  sharp  cutting 
edge,  and  a  holder  (figure  19a,  b),  a  bark  disk  bearing  a  portion  of  a 
lichen  thallus  could  be  removed  from  a  tree  with  little  injury  to  the  lichen. 
The  disk  could  then  be  transferred  to  a  hole  made  in  the  bark  of  any 
other  or  the  same  tree  using  the  same  bark-borer.  The  death  of  the  inner 
tissues  of  the  bark  disk  was  found  to  have  no  noticeable  effect  on  the 
attached  thalli.  The  cut  edges  of  the  lichens  themselves  also  showed  no 
degeneration  and,  in  the  case  of  the  control  disks,  continued  to  grow 
after  transplantation. 

With  continued  use  of  the  bark-borer,  the  blade  tended  to  overlap  at 
the  point  where  the  edges  met  (figure  19b).  This  resulted  in  uneven 
disk  edges  and  occasionally  prevented  an  easy  removal  of  the  disk  from 
the  tree.  To  prevent  this,  a  wooden  disk  was  made  %  inch  thick,  and  cut 
so  that  it  fitted  snugly  on  the  inside  of  the  blade  and  against  the  holder. 
This  disk  effectively  prevented  the  overlapping  of  the  blade  during  the 
cutting  operation  and  still  left  sufficient  room  inside  so  that  no  damage 
to  the  lichen  thallus  occurred. 

Two  methods  of  fastening  the  disk  into  its  new  position  were  tried, 
both  employing  grafting  wax  as  an  adhesive.  The  first  (Brodo,  1961b) 
was  to  apply  the  wax  to  the  back  of  the  disk,  and  the  second  was  to 
apply  the  wax  to  the  inside  of  the  hole  receiving  the  disk.  Due  to  the 
much  larger  number  of  disks  lost  in  the  second  year  run,  it  is  recom¬ 
mended  that  the  former  technique  be  used. 

The  transplant  experiments  were  used  primarily  to  study  vertical 
distribution  and  east-west  distribution  (New  York  City  effects)  and  will 
be  discussed  further  under  those  headings.  In  all  cases,  the  lichens  were 
examined  at  least  twice  after  transplantation,  first,  after  4  months,  and 
second,  after  1  year. 

In  addition  to  the  general  methods  described  above,  certain  special 
techniques  and  procedures  are  discussed  in  their  appropriate  sections 
below.  Results  of  individual  studies  are  also  discussed  within  the  sections. 

SUBSTRATE 

Although  lichen  thalli  have  usually  been  considered  as  neither  sapro¬ 
phytic  nor  parasitic,  it  has  long  been  known  that  certain  lichens  are 
more  or  less  restricted  to  certain  substrate  groups.  Keys  to  crustose  lichens 
almost  always  make  use  of  substrate  early  in  the  separation  of  groups  of 
species  on  a  gross  level,  such  as  the  choice  between  “corticolous”  and 
“saxicolous.”  The  degree  of  substrate  specificity,  particularly  of  cortico¬ 
lous  lichens,  has  been  the  subject  of  several  studies  (Hale,  1955a;  Culber¬ 
son,  1955a;  Barkman,  1958;  Brodo,  1959.  1961a). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


21 


In  an  earlier  study  of  Long  Island  lichens  ( Brodo,  1961a),  eight 
corticolous  species  were  categorized  according  to  their  associations  with 
each  of  three  tree  species,  Pimis  rigida,  Quercus  alba,  and  Q.  velutina 
(including  Q.  coccinea  and  Q.  rubra )  in  four  segments  of  the  pine  to  oak 
forest  continuum.  Various  relationships  were  seen:  (a)  significant  posi¬ 
tive  association  of  the  lichen  with  the  tree  species  over  the  entire  con¬ 
tinuum,  (b)  significant  positive  association  in  some  segments,  but  not  in 
all,  (c)  no  significant  positive  or  negative  association  with  the  tree  in 
any  segment,  (d)  significant  negative  association  in  some  segments  but 
not  in  others,  and  (e)  significant  negative  association  in  all  the  con¬ 
tinuum  segments.  The  above  relationships  were  interpreted  as  follows, 
respectively:  (a)  the  lichen  shows  constant  substrate  specificity  indicat¬ 
ing  possible  substrate  requirements,  (b)  the  lichen  shows  some  specificity 
for  the  tree  but  exhibits  no  clearcut  requirement  for  it,  (c)  the  lichen 
shows  considerable  flexibility  in  substrate  requirements,  varying  in  degree 
of  association  with  any  particular  tree  species  as  the  hark  characteristics 
such  as  texture,  chemistry,  and  moisture  relations  change  in  the  different 
stands,  (d)  the  lichen  shows  some  tolerance  for  the  normally  unfavorable 
substrate,  but  will  occur  more  abundantly  on  other  more  favorable  trees 
if  they  are  available,  and  (e)  the  lichen  has  some  sort  of  physical  or 
physiological  inability  to  inhabit  the  substrate. 

The  results  of  that  study  placed  Parmeliopsis  placorodia  in  category 
a  with  respect  to  Pinas  rigida;  Graphis  scripta  and  Lecanora  caesiorubella 
were  in  category  a  with  Quercus  velutina  and  in  categories  e  and  d, 
respectively,  with  Quercus  alba.  The  other  species,  Parmelia  caperata, 
P.  rudecta,  P.  subaurifera,  and  P.  sulcata  had  little  difference  in  their 
associations  with  the  two  oak  species,  although  all  showed  greater  tend¬ 
encies  toward  positive  association  with  black  oak  (possibly  due  to  bark 
stability).  Thus,  caution  is  necessary  in  interpreting  association  tenden¬ 
cies,  since  association  values  vary  somewhat  between  stands  and  vege¬ 
tation  types  (Brodo,  1959). 

From  field  observation  and  collection  data,  a  number  of  other 
lichens  can  be  considered  narrowly  substrate-specific,  although  in  the 
absence  of  unbiased  sampling  no  quantitative  statement  can  be  made 
concerning  them.  Some  of  these  species  are  listed  below  with  their 
substrate  placed  in  parentheses. 

Corticolous:  Alectoria  nidulifera  (Pinus  rigida),  Cetraria  fendleri 
(Pinus  rigida),  Leptorhaphis  epidermidis  (Betula  populifolia),  Lecidea 
anthracophilia  (Pinus  rigida),  Lecidea  scalaris  (Pinus  rigida),  Trypeth- 
elium  virens  (Ilex  spp.  and  Fagus  grandifolia). 

Saxicolous:  Caloplaca  citrina  (mortar  and  concrete),  C.  feracissima 
(concrete),  C.  flavovirescens  (concrete),  Candelariella  aurella  (con¬ 
crete),  C.  vitellina  (granite),  Lecanora  dispersa  (concrete),  Lecidea 
erratica  (granite  pebbles),  Rhizocarpon  obscuration  (granite),  Rinodina 
oreina  (granite),  Sarcogyne  clavus  (granite). 


22 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Terricolous:  Baeomyces  roseus  (eroded  sandy  loam),  Cladonia  sub- 
mitis  (acid  sand),  C.  boryi  (acid  sand). 

Lignicolous:  Chaenotheca  phaeocephala  (white  cedar  stumps), 
Lecidea  aeruginosa  (planks),  Micarea  prasina  (rotting  wood). 

The  statistical  studies  cited  on  p.  20  attempted  to  clarify  the  basic 
factors  involved  in  specificities  by  relying  on  the  correlation  of  lichen 
presence  with  measurable  bark  characters.  Some  of  these  characters  are 
listed  and  discussed  below. 

1.  Texture:  The  external  texture  of  the  substrate  can  be  important 
in  trapping  diaspores  and  protecting  developing  thalli,  in  providing  en¬ 
trance  to  other  layers  or  tissues  of  the  substrate,  in  capturing  and  retain¬ 
ing  moisture  and  chemical  substances,  and  in  supporting  other  organisms 
which  may  remove  potential  lichen  sites,  or  aid  in  any  of  the  above. 
Different  parts  of  a  tree  trunk  may  have  different  textures  and  conse¬ 
quently  may  bear  entirely  different  lichen  floras.  For  example,  in  fissured 
bark  the  rough  fissures  often  bear  hygrophytic  species  such  as  Lepraria 
incana,  whereas  adjoining  plates  support  only  hardy  species  such  as  the 
alga  Protococcus  viridis  (see  Barkman,  1958,  p.  33).  LeBlanc  (1962) 
cites  bark  moisture  capacity  as  causing  the  differences  between  the  rich 
lichen  flora  of  red  oaks  and  the  poor  flora  of  beeches.  It  is  probable, 
however,  that  the  conspicuous  differences  in  bark  texture  between  the 
two  trees  were  important  in  producing  the  different  epiphytic  vegetations. 

Rough  boulders  normally  bear  more  lichens  than  smooth  ones,  and 
some  lichens  undoubtedly  have  adapted  to  growing  on  very  smooth  sur¬ 
face  in  response  to  the  competitive  advantage  of  such  an  ability  (e.g., 
Lecidea  erratica,  Rhizocarpon  obscuration,  Verrucaria  microspora,  and 
V .  silicicola) . 

2.  Moisture-holding  capacities.  Many  studies  dealing  with  epiphytic 
vegetation,  particularly  cryptogamic  vegetation,  have  included  moisture 
capacity  measurements  of  bark  substances  (Billings  and  Drew,  1938; 
Young,  1938;  Hale,  1955a;  Culberson,  1955a;  Barkman,  1958;  Brodo, 
1959;  LeBlanc,  1962).  Barkman  (1958)  has  reviewed  this  subject  in 
detail. 

Although  the  methods  employed  by  the  various  workers  varied 
somewhat,  in  general  it  was  found  that  moisture  capacities  are  greatest 
with  soft,  flaky  barks,  near  the  tree  bases,  on  windward  sides  of  tree 
trunks,  and  in  humid  areas.  Except  in  a  few  cases,  moisture  capacity  was 
expressed  as  the  ratio  of  water  absorbed  to  dry  weight  of  the  sample. 
LeBlanc  pointed  out  that  by  using  dry  weight  in  the  expression,  barks 
with  the  same  actual  moisture  capacity  per  unit  of  exposed  surface  may 
appear  to  have  different  moisture  capacities  if  their  densities  are  differ¬ 
ent.  For  example,  bark  sample  A  with  a  surface  area  of  10  cm2  and 
weighing  10  grams  may  absorb  5  grams  of  water  when  submerged. 
Sample  B,  also  with  a  surface  area  of  10  cm2  (and  of  the  same  volume) 
but  weighing  20  grams  may  also  absorb  5  grams  of  water.  Since  sample 
B  is  twice  as  dense  (and  weighs  twice  as  much  as  sample  A,  it  appears 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


23 


to  have  on!y  half  the  moisture  capacity,  when  in  reality  the  capacities  are 
equal.  LeBlanc  (1962)  attempted  to  correct  for  this  error  by  expressing 
water  gain  on  a  “per  unit  surface  area”  basis.  Unfortunately,  it  is  ex¬ 
tremely  difficult  to  accurately  measure  surface  area  with  any  but  the 
smoothest  of  bark  types,  and  serious  errors  may  thus  be  introduced  into 
expressions  derived  in  this  way. 

Barkman  ( 1958)  stated  that  moisture  capacity  is  more  meaningful  if 
presented  in  terms  of  sample  volume.  This  would  be  excellent  to  compare 
barks  which  are  known  to  become  totally  and  uniformly  saturated  with 
water.  However,  if  only  surface  layers  are  wetted,  as  might  well  be  the 
case  with  some  of  the  hard-barked  oaks,  moisture  capacity  per  unit 
volume  is  unusable. 

The  measurement  of  the  rate  at  which  a  given  bark  sample  returns 
to  dry  weight  after  being  saturated  (either  by  vapor  or  by  liquid  water) 
for  any  given  period  of  time  is,  under  uniform  conditions  of  humidity, 
a  direct  function  of  its  surface  area,  moisture  capacity,  and  water  bind¬ 
ing  capacity,  all  parameters  of  importance  to  epiphytes.  Hale  (1955a) 
and  Billings  and  Drew  (1938)  presented  some  data  on  water  loss  and 
found  that  bark  samples  returned  to  approximately  dry  weight  in  about 
the  same  time  for  all  trees  studied,  but  that  the  initial  rates  were  greatest 
in  tree  barks  having  the  highest  moisture  capacity  (expressed  as  a  per¬ 
cent  dry  weight).  Although  these  figures  are  important,  they  still  do  not 
reveal  exactly  how  much  water  remains  available  to  the  epiphyte  during 
this  water  loss.  That  is,  a  bark  which  absorbs  four  grams  of  water  per 
unit  surface  area  may  lose  water  twice  as  fast  during  the  first  hour  as  a 
bark  which  absorbs  one  gram  of  water  per  unit  area,  but  at  the  end 
of  that  hour  the  former  still  contains  twice  as  much  water  as  the  latter 
and  in  a  less  strongly  bound  and  hence  more  available  state.  Thus,  it 
would  seem  that  water  loss  rates  alone  could  not  be  used  as  a  substitute 
for  some  sort  of  a  moisture  capacity  expression. 

There  seems  to  be  no  way  to  entirely  overcome  the  density  or  the 
surface  area  problems.  It  is  possible  to  introduce  a  correction  factor 
into  the  dry  weight  expression  to  eliminate  the  density  error,  but  the 
result  is  an  expression  in  terms  of  volume  and,  as  stated  above,  the 
samples  would  then  have  to  be  of  the  same  volume  or  be  proven  to  be¬ 
come  uniformly  saturated  with  water.  The  following  example  will 
illustrate  this  point. 


Sample  A 


Sample  B 


Dry  weight: 
Volume: 

Density: 

Water  absorbed: 
Moisture  capacity: 
Density  correction: 


0.5  gm/cm3 
6  gm 

6  gm/5  gm  =  1 .20 


m.c  X  d  = 

1 .20  X  0.5  =  0.60 


5  gm 

1 0  cm3 


6  gm/3  gm  =  2.00 
m.c  X  d  = 

2.00  X  0.3  =  0.60 


3  gm 
1 0  cm3 
0.3  gm/cm3 
6  gm 


24 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


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LICHENS  OF  LONG  ISLAND,  NEW  YORK 


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26 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Table  lb.  Trees  ranked  in  order  of  their  bark  moisture  capacity.  The 
data  are  from  table  la.  Number  1  has  the  highest  moisture  capacity, 
and  number  7  has  the  lowest. 


Surface 


With  data  from 

Dry  weight 

Volume 

area 

MESIC 

i. 

Ulmus 

Ulmus 

Q.  cocc. 

i 

k 

2 

Q.  alba 

Fagus 

Ulmus 

3. 

Fagus 

Q.  alba 

Q.  vel. 

4. 

Q.  cocc. 

Q.  cocc. 

Q.  rubra 

5. 

Pinus 

Q.  vel. 

Q.  alba 

>| 

y 

6. 

Q .  vel. 

Q.  rubra 

Fagus 

XERIC 

7. 

Q.  rubra 

Pinus 

Pinus 

Table  2.  Sand 

and  soil  pH.  All  samples  were  from  uniform  surface 

material,  mixed  into  a  slurry  with  distilled  water,  equilibrated  for  about 
15  minutes,  and  measured  with  glass-electrode  pH  meter.  Only  one 
sample  from  each  source  was  studied. 


General  category 

Specific  source  locality 

pH 

Exposed,  eroding 

Yaphank 

4.3 

ground,  supporting 
Baeomyces;  central 

Commack 

4.2 

Long  Island 

Riverhead 

4.1 

Beach  sand  from 

Fire  Island  (Bellport) 

4.5 

south  shore 

Napeague  Beach 

4.6 

Sand  from  central 
part  of  island 

near  Manorville 

4.2 

Sand  from  north 

Shoreham,  beach  sand 

6.2 

shore  beaches  and 

behind  very  low  dunes 

dunes 

Rocky  Point,  top  of  high 
bluff,  facing  L.  I.  Sound 

6.1 

Rocky  Point,  over  crest  of 
bluff,  protected  from  full 
on-shore  winds 

5.1 

Rocky  Point,  on  beach 

5.8 

Transplanted 

From:  Fire  Island  (south  shore); 

4.5 

samples  of 

to:  Bellport  on  Great  South 

beach  sand 

Bay;  for  one  year 

From:  Fire  Island  (south  shore); 
to:  Shoreham  (north  shore,  on 
beach  behind  very  low  dunes); 
for  one  year. 

5.8 

LICHENS  OF  LONG  ISLAND,  NEW  YORK 


27 


These  final  values  represent  grams  water  absorbed  per  unit  volume, 
and  are  functions  of  moisture  capacity  which  are  comparable  and  give 
relative  positions  of  the  bark  types.  The  principle  is  valid,  hut  is  replete 
with  difficult  problems,  some  of  which  were  mentioned  above.  In  addi¬ 
tion,  sample  volumes  are  almost  impossible  to  keep  constant,  since  some 
bark  types  are  thin  (e.g.,  Fagus  granclifolia,  Acer  rubrum)  and  others 
must  be  taken  in  thick  slices  (e.g.,  Quercus  rubra,  Ulmus  americana) . 
The  measurement  of  volume  is  somewhat  inaccurate  in  certain  bark 
types,  since  bark  samples  often  contain  spaces  which  trap  air  in  water 
displacement  procedures.  Inaccurate  volume  determinations,  of  course, 
make  density  figures  of  little  practical  value.  Errors  in  volume  measure¬ 
ment  also  decrease  the  value  of  moisture  capacity  expressions  based 
on  volume  alone. 

Despite  the  shortcomings  of  some  of  the  methods  discussed  above, 
moisture  capacity  measurements  were  performed  on  bark  samples  from 
a  number  of  common  Long  Island  trees.  The  methods  employed  were 
essentially  those  of  Culberson  (  1955a). 

Bark  samples  were  obtained  with  the  use  of  the  bark-borer  used  in 
the  transplant  studies  ( p.  19-20)  wherever  possible.  This  method  provided 
samples  of  very  similar  size  and  volume  except  for  the  thin  bark  trees. 
Some  bark  types  were  not  amenable  to  bark  disk  removal  due  to  their 
instability  and  flaky  nature  (e.g.,  Quercus  alba  and  Pinus  rigida) .  Bark 
samples  of  these  trees  were  collected  without  a  borer  and  were  cut 
down  to  approximate  the  surface  area  of  the  disks. 

The  samples  remained  unstudied  for  a  year  and  a  half  and  so  were 
quite  dry  at  the  beginning  of  the  observations.  They  were  oven  dried  at 
100°  C.  for  a  period  of  20  hours  to  ensure  uniform  dessication.  After 
cooling,  the  samples  were  weighed,  coated  with  a  layer  of  paraffin  on 
all  cut  surfaces,  and  then  reweighed  to  derive  the  weight  of  the  wax.  The 
volume  of  the  wax  on  each  sample  was  calculated  with  a  knowledge  of 
the  wax’s  density. 

Volume  was  measured  by  water  displacement  and  was  precise  to 
0.3  cc  but  volume  measurements  were  somewhat  exaggerated  in  certain 
bark  types  having  large  amounts  of  air  retention,  e.g.,  Pinus  rigida  bark. 
The  exposed  area  was  measured  as  follows:  a  small  piece  of  aluminum 
foil  was  carefully  fitted  to  the  contours  of  a  bark  sample;  the  excess 
foil  was  then  cut  off  at  the  limit  of  the  exposed  colonizable  surface;  the 
fitted  foil  piece  was  pressed  flat,  numbered,  and  weighed;  the  weights 
of  the  various  foil  replicates  were  then  fitted  on  a  standard  curve  con¬ 
structed  from  the  weights  of  foil  samples  of  known  surface  area  to  find 
the  surface  areas  of  the  bark  samples.  In  this  way  very  irregular,  rough 
surface  features  of  the  bark  samples  could  be  accounted  for  in  the  surface 
area  measurements. 

Water  absorption  expressed  as  per  unit  dry  weight,  per  unit  volume, 
and  per  unit  of  colonizable  surface  is  presented  in  table  I  along  with 
the  other  hark  features  of  the  common  Long  Island  trees. 


28 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


In  ranking  the  trees  in  order  of  bark  moisture  capacity,  we  can 
see  that  the  method  of  expression  is  very  important  in  the  relative  posi¬ 
tions  of  the  various  species.  Dry  weight  and  volume  expressions  matched 
most  closely,  with  Pinus  rigida  being  a  notable  exception.  Volume  meas¬ 
urements  of  pine  bark  are  complicated  by  considerable  air  retention 
between  the  bark  plates  during  water  displacement  as  noted  above. 
This  error  would  make  the  volume  appear  larger  than  it  actually  is 
and  would  thus  effectively  "lower”  the  moisture  capacity  expression 
based  on  volume.  Quercus  rubra  appears  more  mesic  than  Fagus  in  the 
surface  area  expression,  whereas  the  opposite  is  true  with  the  dry  weight 
and  volume  expressions.  LeBlanc  (1962)  noted  the  same  change  in  rela¬ 
tive  position  of  the  two  trees  in  his  studies.  One  important  difference  in 
the  area,  dry  weight,  and  volume  sequences  is  in  the  relative  position  of 
Quercus  alba;  it  appears  relatively  more  xeric  in  the  first  and  more  mesic 
in  the  latter  two.  In  view  of  the  strong  emphasis  which  has  been  placed 
on  the  difference  in  moisture  capacities  between  black  and  white  oaks  in 
the  past  (see  Hale,  1955a),  it  may  be  well  to  recheck  these  findings 
in  other  areas  with  larger  samples.  The  sample  size  (1  to  8)  in  the  data 
presented  here  was  too  small  to  warrant  the  formulation  of  strong  con¬ 
clusions  pertaining  to  the  relative  positions  of  various  trees  with  regard 
to  their  bark  types. 

3.  Stability.  The  rapidity  with  which  a  given  substrate  surface  is 
removed  or  changed  in  some  way  has  a  strong  influence  in  determining 
which  lichens  will  inhabit  the  surface.  Only  rapidly  growing  and  matur¬ 
ing  species  can  become  established  on  unstable  surfaces. 

No  species  can  colonize  shifting  sand  as  is  found  on  sand  dunes. 
Dune  species  usually  become  established  on  relatively  less  active  dunes, 
on  plant  remains  ( Brodo,  1961a),  or  under  the  protection  of  trailing  or 
low  growing  vascular  species  such  as  Arctostaphylos  uva-ursi  or  Hud- 
sonia  tomentosa.  The  thalli  may  later  become  detached  and  continue 
development  independently  on  the  relatively  stable  sand  surface.  Where 
the  sand  is  protected  from  strong  wind  action  and  becomes  covered  with 
an  organic  film,  as  in  scrub  oak  thickets,  certain  species  such  as  Lecidea 
uliginosa,  L.  granulosa,  or  Cladonia  cristatella  can  become  established 
and  actually  serve  in  binding  the  sand  particles  together  (p.  42).  Where 
the  sand  is  even  more  stabilized,  many  more  terrestrial  species  may  gain 
foothold.  Baeomyces  roseus  and  Pycnothelia  papillaria  can  apparently 
grow  fast  enough  to  grow  over  the  eroding  surfaces. 

Rapidly  sloughing  bark  severely  limits  the  number  of  species  which 
can  inhabit  a  tree  (Hale,  1952a;  Barkman,  1958),  and  it  is  likely  that 
this  is  one  of  the  reasons  for  the  relatively  small  number  of  species 
found  on  Pinus  rigida.  The  best  development  of  any  species  growing  on 
pine  occurs  on  the  edges  of  the  bark  plates  deep  in  the  fissures  where  the 
bark  is  most  stable.  The  poorest  development  is  on  the  plate  surfaces  which 
lose  outer  flakes  of  bark  almost  continuously.  The  role  of  bark  stability 
in  limiting  species  coverage  is  made  strikingly  clear  when  the  standing 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


29 


trunk  of  a  dead  pine  appears  close  to  a  living  tree.  The  stable  bark  of  a 
dead  tree  is  covered  with  lichens,  whereas  only  spotty  coverage  is  seen 
on  the  living  ho'e,  even  though  both  trunks  have  equal  light  and  are 
standing  side  by  side.  It  is  possible  that  the  absence  of  a  canopy  may 
have  some  effect  in  changing  the  moisture  relations  (via  increasing  drain¬ 
age)  on  the  dead  tree  or  in  failing  to  contribute  inhibitory  organic  ma¬ 
terial,  but  there  are  probably  not  as  important  as  the  stabilized  substrate. 

Pebbles  and  small  stones  often  shift  and  roll  with  changes  in 
weather,  and  thereby  expose  or  cover  lichens  which  may  be  growing  on 
their  surfaces.  I  first  considered  this  problem  in  a  study  of  the  lichens 
of  an  old  field  at  the  American  Museum  of  Natural  History  Biological 
Laboratory  at  Dix  Hills,  Long  Island.  Yearly  observations  of  numerous 
pebbles  indicated  that  Lecidea  erratica  develops  very  quickly  on  exposed 
stones  (p.  42  to  43).  Since  these  pebbles  and  stones  undoubtedly  shift  or 
even  turn  over  with  frost  action  and  heavy  rains,  rapid  growth  may  be 
an  important  factor  in  the  maintenance  of  populations  of  these  species. 
Typical  members  of  the  exposed  boulder  communities  such  as  Parmelia 
arseneana  and  P.  conspersa  have  been  found  on  some  stabilized  pebbles, 
adding  strength  to  the  supposition  that  the  instability  of  pebbles  may  be  a 
factor  in  eliminating  these  overshadowing  but  slowly  growing  species 
from  competing  for  space  with  the  small  but  rapidly  developing  Lecideae. 

Small  stones  continuously  roll  and  shift  in  the  littoral  zone  of  the 
shallow  bays  and  inlets,  and  it  is  not  surprising  to  find  that  the  marine 
Verrucariae  (V.  microspora  and  V.  silicicola )  often  are  found  growing  on 
all  sides  of  these  pebbles,  regardless  of  their  position  when  collected. 

4.  Chemical  composition.  Bark  chemistry  as  with  bark  moisture 
capacity  has  been  studied  by  most  epiphyte  ecologists.  Barkman  (1958) 
again  provides  an  excellent  summary  of  the  information  published  on 
the  subject. 

Of  the  many  facets  of  bark  chemistry,  acidity  has  been  the  most 
widely  studied.  Great  emphasis  has  been  placed  on  bark  acidity  in  ex¬ 
plaining  the  distribution  of  some  lichens  (Billings  and  Drew,  1938;  Hale, 
1955a;  Culberson,  1955a;  Barkman,  1958;  DuRietz,  1945  in  Almborn, 
1948).  Barkman  (1958)  and  Almborn  (1948)  have  pointed  out  some  of 
the  oversimplifications  to  which  some  authors  have  fallen  victim,  but 
pH  remains  an  important  factor  to  be  considered  in  epiphytic  ecology. 

The  pH  of  bark  samples  of  several  of  the  common  trees  were 
measured.  A  few  grams  of  bark  material  were  obtained  by  slicing  the  sur¬ 
face  layers  from  a  bark  sample  and  chopping  them  into  a  mealy  con¬ 
sistency.  Between  five  and  seven  ml  of  distilled  water  were  added  to 
each  chopped  bark  sample,  enough  to  form  a  thick  slurry,  and  the 
mixture  was  allowed  to  equilibrate  at  room  temperature  (approximately 
5  hours).  Acidity  was  measured  by  using  a  glass  electrode  Beckman 
pH  meter. 

The  acidity  of  soil  samples  was  measured  in  a  similar  way.  A  soil 
slurry  was  formed  using  one  part  water  and  two  parts  soil  (approximately 


30 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


20  cc  soil  and  10  ml  distilled  water).  The  mixture  was  allowed  to 
equilibrate  (15  minutes)  and  pH  was  measured  using  the  same  apparatus 
as  mentioned  above.  (The  results  of  these  measurements  are  given  in 
tables  la  and  2.) 

It  would  seem  that  acidity  either  affects  the  lichen  vegetation  directly 
or  indirectly,  or  reflects  a  condition  which  does,  because  definite  cor¬ 
relations  can  be  seen  between  lichen  presence  and  substrate  pH.  The 
very  low  pH  of  Finns  rigida  bark  could  explain  its  poor  and  restricted 
flora,  and  the  high  pH  of  Ulmus,  and  Robinia  provide  clear  associations 
with  the  so-called  “nitrophytic”  (Xanthorion)  community.  It  is  especi¬ 
ally  significant  that  a  black  oak  once  found  bearing  Xanthoria  thalli 
had  neutral  bark,  although  this  species  of  tree  normally  has  very  acid 
bark.  That  particular  oak  was  growing  in  the  center  of  a  large  Long 
Island  duck  farm,  the  atmosphere  of  which  was  very  obviously  filled 
with  ammonia  and  other  gaseous  and  fine  particulate  materials.  Trees 
along  farm  roads  exposed  to  farm  dust  have  long  been  known  to  bear 
rich  “coniophilous”  communities  ( Barkman.  1958;  Almborn,  1948). 
The  very  high  moisture  capacity  of  Ulmus  may  be  a  significant  factor  in 
the  specificity  of  roadside  species,  although  the  oak  mentioned  above, 
which  supported  a  rich  Xanthoria  community,  had  a  low  moisture  capac¬ 
ity  comparable  with  other  oaks.  The  problem  of  separating  nitrogen 
concentration  from  acidity  in  correlations  of  this  kind  has  been  dis¬ 
cussed  by  both  Almborn  (1948)  and  Barkman  (1958).  Both  authors 
point  out  the  possibility  of  other  factors  being  involved,  especially  phos¬ 
phorous  concentrations.  For  example,  such  typically  “nitrophilous”  spe¬ 
cies  as  Caloplaca  cerina,  C.  pyracea,  and  C.  flavovirescens  are  also  found 
on  turtle  shell  and  bone,  substrates  known  to  be  high  in  phosphorus. 
Since  calcium  concentrations  are  often  high  in  alkaline  substrates,  cal¬ 
cium  may  be  important  in  these  specificities  as  well.  Many  so-called 
“nitrophilous”  lichens,  especially  Xanthoria  parietina ,  X.  fallax,  and 
Physcia  adscendens  are  commonly  found  on  mortar  and  concrete  which 
have  a  high  pH  and  calcium  concentration,  but  are  certainly  not  rich 
in  nitrogen  compounds. 

The  presence  of  Cladonia  submitis  and  associated  lichens  on  the 
south  shore  and  inland  and  their  absence  on  the  north  shore  is  strongly 
correlated  with  soil  acidity.  The  south  shore  and  inland  sands  are  all 
distinctly  acid,  whereas  the  north  shore  sands  are  neutral  (table  2). 
Exactly  what  is  involved  in  this  correlation  is  still  not  clear  ( p.  203). 
The  eroding  soil  supporting  Baeomyces  roseus  has  the  same  pH  as  the 
dune  sand  and,  therefore,  acidity  cannot  explain  the  differences  in  the 
terrestrial  communities  of  the  two  soil  types.  The  higher  moisture  capac¬ 
ity  and  organic  content  of  the  eroded  sandy  loam  are  possibly  the  decid¬ 
ing  factors  in  this  case. 

Other  substrate  minerals  not  studied  here  are  undoubtedly  important 
in  lichen  distributions.  Although  some  data  are  available  on  the  mineral 
contents  of  substrates  (Barkman,  1958)  and  mineral  nutrition  of  lichens 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


31 


(Smith,  1960a,  1960b,  etc.)  the  subject  is  still  far  from  adequately 
understood.  The  extreme  specificity  of  Trypethelium  virens  for  several 
species  of  Ilex  (Johnson,  1959)  suggests  the  presence  of  some  genetically 
controlled  metabolite  in  that  genus  which  is  essential  for  the  establish¬ 
ment  or  survival  of  the  lichen.  Fagus  grandifolia ,  another  common  phoro- 
phyte  for  Trypethelium  virens,  would  then  also  have  to  possess  the 
ability  to  produce  this  substance  or  a  substitute.  There  is  some  evidence 
that  Trypethelium  actually  does  utilize  some  bark  material  (Johnson, 
1940).  Fink  (1913)  suggested  that  other  endophloedal  crustose  species 
also  derive  some  nutritional  benefit  from  their  substrate.  The  fact  that 
Trypethelium  virens  has  been  found  in  healthy  condition  on  Long  Island 
only  on  living  trees,  an  observation  also  made  by  Johnson  (1940),  and 
the  fact  that  all  the  host  trees  of  this  species  have  thin  living  bark,  adds 
weight  to  the  possibility  that  a  specific  class  of  metabolite  is  involved. 

It  is  easy  to  imagine  a  lichen  living  on  a  nutrient-rich  substrate 
making  use  of  these  nutrients,  especially  when  all  the  mechanisms  for 
their  absorption  are  available  and  efficient  (Smith,  1962).  More  work  on 
substrate  specificity  is  needed  to  clear  up  these  important  problems. 

CLIMATE 

Atmospheric  humidity  is  involved  in  the  water  budget  of  a  lichen 
thallus  to  a  greater  degree  than  it  is  in  the  water  budget  of  a  rooted 
vascular  plant  in  the  same  general  habitat.  This  is  due  to  a  lichen’s 
ability  to  pick  up  water  vapor  and  use  the  absorbed  moisture  in  photo¬ 
synthesis  and  metabolism  in  a  relatively  short  period  of  time,  as  com¬ 
pared  with  the  green  parts  of  vascular  plants  (p.  32).  Thus,  a  habitat 
which  might  be  dry  for  a  terrestrial  vascular  plant  due  to  excessive 
soil  drainage,  may  not  be  dry  to  lichens  if  air  humidity  is  high 
enough  during  part  of  the  24  hour  cycle.  It  is  the  microclimate  which 
one  must  measure  in  order  to  characterize  the  water  budget  in  the 
ecological  niche  of  a  lichen.  Unfortunately,  to  perform  such  measure¬ 
ments  was  beyond  the  scope  of  this  work,  although  such  studies  would 
be  extremely  interesting  and  valuable. 

Vertical  and  horizontal  zonation  ( p.  34  to  35)  and  patterned  dis¬ 
tribution  in  bark  fissures  or  on  bark  ridges  are  probably  at  least  partly 
manifestations  of  different  microclimates. 

1.  Illumination  and  temperature.  Light  intensity  is  a  very  complex 
factor,  having  both  direct  and  indirect  effects  on  microclimate.  As  Bark- 
man  (1958,  p.  57)  states  “.  .  .  it  is  often  difficult  to  decide  whether  a 
given  species  is  photophilous,  thermophilous,  or  xerophilous,”  since  strong 
light  will  raise  the  temperatures  of  both  bark  surfaces  and  the  lichens 
themselves  (especially  if  they  are  dark  colored)  and  will,  therefore,  in¬ 
crease  the  evaporation  rate,  which  in  turn  will  increase  drought  condi¬ 
tions.  The  role  of  illumination  in  raising  temperatures,  and  thus  evapora¬ 
tion  rates  and  drought,  was  an  important  consideration  in  Barkman’s 


32 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


(1958)  summary  of  the  causes  of  horizontal  zonation  (zonation  accord¬ 
ing  to  direction  of  exposure)  of  epiphytes  on  tree  trunks  in  Holland. 

Lichens  derive  their  principal  nutrition  from  the  photosynthesis 
products  of  their  algal  components,  and  so  the  lichen  thallus  is  depend¬ 
ent  upon  light  for  survival.  Since,  in  the  lichens  that  have  been  studied, 
the  net  rates  of  photosynthesis  per  unit  surface  area  are  much  lower 
than  those  of  the  leaves  of  higher  plants  (Smith,  1962),  it  is  not  surpris¬ 
ing  that  most  lichens  are  found  in  moderately  or  well-lighted  habitats. 
Deeply  shaded  forests,  dry  or  moist,  are,  in  general,  lichen  poor.  Lichens 
exposed  to  full  sunlight,  however,  are  often  subject  to  extreme  drought. 
Many  species  have  developed  adaptations,  such  as  cortical  pigment 
accumulation  and  cortical  thickenings  (Barkman,  1958)  which  cut  down 
light  intensity  and  transpiration. 

Some  Long  Island  lichens  which  seem  to  be  distinctly  photophilous 
are  Cetraria  islandica  subsp.  crispa,  Cladonia  submitis,  C.  boryi,  Xan- 
thoria  parietina,  Parmelia  sulcata,  Usnea  strigosa,  Ramalina  fastigiata, 
Pertusaria  xanthodes,  and  Lecanora  caesiorubella  subsp.  lathamii.  These 
species  are  most  often  found  in  well-illuminated  habitats  even  though 
their  general  substrate  types  extend  into  more  shaded  areas.  The  first 
three  species  mentioned  above  are  found  almost  exclusively  on  exposed 
sand  plains  and  downs.  Xanthoria  parietina  has  long  been  known  to  be 
photophilous  (Barkman,  1958).  The  remaining  species  occur  most  fre¬ 
quently  in  well-lighted  but  dry,  mature  pine-oak  forests  of  central  Long 
Island.  This  vegetation  type  can  be  thought  of  as  a  compromise  habitat 
between  optimum  light  and  optimum  moisture  (Brodo,  1961a).  In 
more  humid  localities,  such  as  the  Montauk  region  on  the  southern  fluke 
of  Long  Island  (figure  5),  these  species  all  reach  their  maximum  devel¬ 
opment  in  completely  exposed  situations. 

2.  Moisture.  Lichens  are  classically  thought  of  as  among  the  most 
drought-resistant  plant  types.  Although  it  is  true  that  many  species  can 
survive  in  habitats  much  too  dry  to  support  any  but  the  most  xeric  of 
bryophytes,  many  lichens  are  clearly  limited  to  rather  moist  environ¬ 
ments  and  many  others  are  very  sensitive  to  changes  in  environmental 
moisture. 

The  role  of  moisture  in  the  photosynthetic  efficiency  of  lichens  has 
been  reviewed  and  summarized  by  Smith  (1962).  He  points  out  that 
in  the  nonaquatic  lichens  which  have  been  studied,  photosynthetic  effi¬ 
ciency  is  greatest  at  moisture  contents  below  saturation.  In  nature,  non¬ 
aquatic  lichens  are  rarely  saturated.  Although  most  nonsorediose  lichens 
absorb  liquid  water  rapidly,  they  lose  water  almost  as  fast.  Absorption  of 
water  vapor  is  a  much  slower  process,  but  constantly  humid  areas  are 
undoubtedly  less  droughty  than  dry,  exposed  areas  with  frequent  rains 
(Barkman,  1958).  This  is  especially  the  case  since  it  has  been  shown  that 
lichens  can  absorb  water  from  nonsaturated  air  (Pavillard,  1939;  Bark¬ 
man,  1958).  Thus,  the  misty  thickets  and  shrubby  groves  of  the  depre- 
sions  in  the  Montauk  area  are  wet  habitats,  whereas  just  outside  these 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


33 


groves  on  the  exposed  dunes,  where  constant  strong  winds  make  evapo¬ 
ration  high,  the  habitat  is  extremely  dry  (Taylor,  1923;  see  also  Salisbury, 
1952).  Ried  (1960)  pointed  out  that  lichens  are  most  seriously  damaged 
when  they  are  subjected  to  intermittent  wet  and  dry  periods.  This  may 
explain  why  some  lichens  that  were  thought  to  be  drought  resistant 
through  laboratory  experimentation  actually  appear  sensitive  to  low 
moisture  conditions  when  observed  in  the  field.  He  suggested  that  it  is 
the  ability  of  various  species  to  recover  from  a  drought  which  might 
determine  their  distribution. 

Moisture  also  has  an  important  indirect  influence  on  lichen  growth. 
Inasmuch  as  microbial  activity  is  highly  dependent  on  moisture  levels  of 
various  habitats,  any  lichen  distribution  dependent  on  the  products  of 
either  fungal  or  bacterial  growth  or  on  the  changes  in  the  physical 
characters  of  substrates  subjected  to  such  activity  would  necessarily 
follow  moisture  changes  as  well. 

Moisture  comes  to  the  corticolous  lichen  thallus  from  precipitation 
and  from  air  humidity  (both  directly  and  through  the  wetted  substratum), 
and  rarely  by  inundation.  In  the  tropics,  moisture  may  be  made  available 
directly  from  the  living  tissue  of  the  thin-barked  trees  (Imshaug,  pers. 
comm.).  The  evaporation  rate  in  any  particular  habitat  and  the  moisture 
capacity  of  the  substrate  determines  how  efficiently  and  for  how  long 
this  moisture  is  available  to  the  lichen. 

The  availability  of  rain  to  epiphytic  lichens  is  influenced  by  canopy 
type  and  canopy  density  mainly  through  their  effect  on  the  flow  of  water 
from  the  leaves  and  twigs  down  the  branchlets  and  branches  and  finally 
down  the  trunk.  This  flow  of  water  (“stemflow”)  is  often  a  major  route 
for  the  entrance  of  moisture  to  the  forest  interior  (Kittredge,  1948)  and 
is,  of  course,  of  major  importance  to  corticolous  plants.  Stemflow  is 
greatest  with  trees  having  ascending  branches  (“centripetal  crown”)  as  in 
Acer  and  Feigns,  and  is  least  with  trees  having  drooping  branches  (“cen¬ 
trifugal  crown”)  as  in  Picea;  Quercus  and  Pinus  are  intermediate  in  this 
respect  (Barkman,  1958;  Geiger,  1965).  It  should  also  be  borne  in 
mind  that  precipitation  which  has  passed  through  a  canopy  (“through- 
fall”)  is  much  richer  in  certain  minerals  and  ions  than  unintercepted 
rain  (Tamm,  1951). 

In  the  pine-oak  forest  of  Long  Island,  much  of  the  rain  reaches 
the  hole  directly  through  the  loose  canopy  as  well  as  by  stemflow.  In  the 
dense,  red  oak  forest,  light  rains  never  reach  the  ground  or  tree  trunks, 
being  evaporated  directly  from  the  canopy.  Heavy  rains  filter  down 
through  the  canopy,  but  only  reach  the  bole  via  rain  tracks  (the  channels 
of  most  liquid  stem  runoff)  and  general  stemflow.  However,  once  the 
rain  has  wet  the  ground  and  bark  in  a  shaded  forest,  the  precipitation 
is  slowly  converted  to  increased  air  humidity  which  slows  evaporation 
from  the  wetted  thalli  and  supplies  additional  moisture  for  a  long  period. 
The  rain  in  a  pine-oak  forest,  on  the  other  hand,  is  quickly  lost  in  the 
very  well  drained,  sandy  soils  and  dried  from  the  bark  with  no  sub- 


34 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


stantial  increase  in  the  local  humidity  for  more  than  a  very  short  period 
of  time. 

It  is,  therefore,  in  the  relatively  open  habitats  that  hygrophilous 
species  occupy  substrates  with  high  moisture  capacities  (p.  51  to  54). 

VERTICAL  DISTRIBUTION 

The  vertical  zonation  of  corticolous  epiphytes  has  intrigued  many 
cryptogamic  ecologists  (Plitt,  1924;  Billings  and  Drew,  1938;  Hale, 
1952a;  Culberson,  1955a;  Barkman,  1958;  Brodo,  1959,  1961a,  1961b). 
Methods  of  study  varied  from  detailed  investigations  of  a  few  trees  from 
base  to  crown  (Plitt,  1924;  Hale,  1952a)  to  studies  of  hundreds  of 
trees  only  at  basal  and  breast  height  quadrats  (Hale,  1955a;  Culberson, 
1955a;  Brodo,  1961a).  Barkman  (1958)  made  numerous  observations 
concerning  vertical  zonation  and  thoroughly  reviewed  the  previous  work. 

Several  approaches  were  taken  in  the  study  of  this  phenomenon 
on  Long  Island:  a  statistical  evaluation  of  species  presence  in  breast 
height  and  basal  quadrats,  experimental  transplant  studies,  and  field  obser¬ 
vations  of  lichen  communities. 

As  a  result  of  the  statistical  investigations  described  previously, 
several  common  species  could  be  characterized  as  to  their  vertical  zona¬ 
tion  affinities  (table  3  and  Brodo,  1961a.  From  an  examination  of  the 
vertical  distribution  of  certain  common  corticolous  lichens  in  the  pine- 
oak  forests,  as  compared  with  the  red  oak  forests  (disregarding  phoro- 
phyte  species)  (table  3),  one  can  see  that  the  frequencies  in  the  basal 
quadrat  in  the  former  are  consistently  higher  than  those  in  the  latter. 
This  tendency  of  species  normally  dwelling  at  breast  height  to  be  con¬ 
fined  to  the  basal  area  in  dry  pine  oak  woods  is  consistent  with  the  state¬ 
ments  made  by  several  authors  (Billings  and  Drew,  1938;  Plitt,  1924; 
Potzger,  1939;  Barkman,  1958)  concerning  vertical  microclimatic  gradi¬ 
ents.  That  is,  bark  moisture  is  greater  and  evaporation  is  slower  at  tree 
bases,  as  opposed  to  microhabitats  higher  on  the  trunk.  Barkman  (1958) 
has  pointed  out  how  different  moisture  conditions  in  different  vegetation 
types  can  influence  epiphytic  vertical  distributions.  He  states  that  in 
moist  woods  typically  base-dwelling  communities  sometimes  cover  entire 
trunks.  On  Long  Island,  this  phenomenon  is  particularly  striking  with 
bog  tree  epiphytes.  Within  the  humid,  cool  bogs,  Lobaria  pulmonaria  and 
Lobaria  quercizans  grow  at  all  levels,  but  just  outside  the  bogs,  in  the 
drier  oak  forests,  the  same  lichens  are  confined  to  tree  bases. 

Transplant  experiments  done  in  1960-61  concerned  with  vertical 
distribution  of  Lecanora  caesiorubella  and  Cladonia  chlorophaea  ( Brodo, 
1961b)  showed  that  the  Lecanora  could  survive  when  transplanted  from 
breast  height  to  the  tree  base,  but  that  the  Cladonia ,  upon  being  trans¬ 
ferred  from  the  base  to  breast  height  (1.3m),  soon  decayed.  Lichen- 
frequency  data  bear  out  the  supposition  that  the  Lecanora  is  somewhat 
more  facultative  in  its  vertical  distribution  that  is  the  Cladonia  (table  3). 
Since  the  lichens  were  transferred  on  their  original  intact  substrate  to 


35 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 

Table  3.  Vertical  distribution  of  some  corticolous  lichens  in  red  oak 
and  pine-oak  forests.  Not  all  species  listed  were  treated  in  the  pine-oak 
forest  data  because  some  were  absent  or  too  infrequent,  and  some,  due  to 
recognition  problems  with  sterile  material,  were  not  included  in  early 
sampling.  Red  oak  forest  data  were  collected  from  localities  7  to  1 1  in 
the  north  shore  transect.  The  pine-oak  forest  data  are  from  continuum 
segments  A  and  B,  in  Brodo  (1961a). 

RED  OAK  FOREST  PINE-OAK  FOREST 


Total 
frequency 
(percent  of 
300  trees) 

Percent 
of  total 
quadrat 
occurrences. 
Base  1.3  m. 

Total 
frequency 
(percent  of 
300  trees) 

Percent 
of  total 
quadrat 
occurrences. 
Base  1.3  m. 

Cladonia  chlorophaea 

9 

100 

0 

42 

96 

4 

C.  coniocraea 

62 

93 

7 

G  rap  his  scrip  ta 

10 

22 

78 

Hypogymnia  physodes 

3 

37 

63 

Lecanora 

caesiorubelia 

6 

6 

94 

L.  chlarotera 

5 

25 

75 

1  1 

33 

67 

Parmelia  caperata 

24 

76 

24 

4 

67 

33 

P.  rudecta 

24 

69 

31 

3 

86 

14 

P.  saxatilis 

14 

57 

43 

11 

60 

40 

P.  sulcata 

22 

9 

91 

45 

21 

79 

Pertusaria  xanthodes 

4 

38 

62 

Physcia  millegrana 

3 

40 

60 

8 

16 

84 

Ph.  orbicularis 

3 

100 

0 

5 

71 

29 

points  on  the  same  tree,  the  experiments  also  strongly  indicated  that  it 
is  microclimatic  conditions,  rather  than  bark  surface  features  or  differ¬ 
ences  in  organic  or  inorganic  nutrients  on  a  vertical  gradient,  which 
largely  determine  where  on  a  particular  tree  a  lichen  can  survive.  Since 
the  degree  of  fungal-bacterial  breakdown  of  bark  appears  to  increase 
towards  the  tree  base,  it  is  possible  that  the  microclimatic  gradient  may 
be  operating  through  a  biological  link  to  influence  the  lichen. 

In  various  local  habitats  not  sampled  in  the  statistical  work,  some 
noteworthy  types  of  vertical  zoning  were  observed.  On  the  windward 
sides  of  trees  growing  close  to  bays  and  lakes,  basal  lichen  communities 
often  extend  far  up  the  trunk  (see  also  Billings  and  Drew,  1938;  Bark- 
man,  1958).  For  example,  Parmelia  nidecta,  Parmelia  caperata,  and 
Physcia  orbicularis,  all  dominantly  base-dwelling  under  normal  condi¬ 
tions,  were  found  growing  high  on  the  bay-facing  side  of  an  oak  tree 
on  Shelter  Island.  The  lee  side  of  the  trunk  had  a  normal  basal  zone. 

Inclination  of  the  phorophyte  trunk  greatly  changes  its  moisture 
conditions  and  permits  basal  vegetation  to  grow  much  farther  towards 
the  crown  (Barkman,  1958).  A  tree  growing  on  a  steep  hillside  essen¬ 
tially  has  the  ground  brought  closer  to  the  crown  on  the  uphill  side  of 
the  trunk,  and  this  side,  then,  has  a  more  “basal”  epiphytic  flora. 


36 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Crustose  species  are  almost  entirely  confined  to  areas  above  the  base, 
except  for  the  normally  basal  epibryic  and  leprose  crusts. 

Although  light  has  classically  been  cited  as  one  of  the  main  causes 
of  vertical  zonation  of  epiphytes  (Plitt  and  Pessin,  1924;  Barkman,  1958; 
Hale,  1952a,  1955a;  etc.),  light  probably  was  not  significantly  involved 
in  the  results  of  the  Long  Island  studies,  since  illumination  does  not 
seem  to  be  a  controlling  factor  in  either  forest  type  at  the  basal  or  1.3m 
levels.  In  pine-oak  forests,  light  appears  to  be  abundant  and  uniform  over 
most  of  the  tree  due  to  the  low,  loose  canopy.  In  the  shaded  red  oak 
forest,  light  appears  to  be  uniformly  low  until  one  reaches  the  upper 
portions  of  the  trunk  and  canopy  far  above  the  level  examined.  On  the 
few  felled  or  windblown  trees  that  were  examined,  an  obvious  trend 
toward  a  greater  number  of  crustose  species  at  the  treetops  and  a  greater 
lichen  cover  in  general  points  to  a  light  effect.  That  photophilous  Usnea 
strigosa  is  most  abundant  in  forest  glades,  forest  edges,  and  on  treetops 
makes  evident  the  fact  that  light  is  the  controlling  force  in  its  vertical 
distribution. 

Moisture,  of  course,  is  of  major  importance  in  all  types  of  lichen 
distributions  (p.  32-34).  Vertical  moisture  gradients  of  many  kinds 
have  been  reported,  including  evaporation  rate  (Plitt  and  Pessin,  1924; 
Potzger,  1939),  bark  moisture  (Billings  and  Drew,  1938;  Hale,  1952a), 
and  relative  humidity  (Barkman,  1958).  With  the  ground  being  a  major 
water  reservoir,  it  is  evident  that  the  farther  one  moves  away  from  this 
reservoir,  the  drier  the  microclimate  will  be.  The  more  humid  an  area, 
the  less  will  be  the  difference  between  humidity  at  ground  level  and 
humidity  at  greater  heights  and,  therefore,  the  less  pronounced  will  be 
the  vertical  vegetational  zonation  which  responds  to  this  moisture  gradi¬ 
ent  (Barkman,  1958,  p.  39).  This  is  indeed  what  is  observed  in  the  Long 
Island  studies,  and  we  can  therefore  conclude  that  moisture  is  probably 
a  controlling  factor  in  most  cases. 

Temperature  and  bark  characteristics  such  as  color,  hardness,  and 
porosity  all  have  their  effects  on  substrate  moisture  relations  via  evapora¬ 
tion  rate  or  moisture  capacities,  and  all  show  vertical  changes.  Since 
epiphytes  are  sensitive  to  moisture  changes,  it  is  easily  seen  how  a  vertical 
zonation  of  epiphytes  can  be  influenced  by  these  physical  features  of 
the  substrate. 

Organic  and  inorganic  nutrients,  having  been  either  blown  on  to  the 
bark  surface,  carried  down  by  stemflow  or  throughfall  (Tamm,  1951),  or 
produced  there  by  local  microbial  activity,  are  distributed  along  a  verti¬ 
cal  gradient  and  may  play  an  important  part  in  the  distribution  of  certain 
species,  particularly  those  that  normally  grow  on  the  ground.  The  possible 
role  of  nutrient  accumulation  in  the  maintenance  of  established  colonies 
of  Cladonia  chlorophaea  has  been  disproven  by  transplant  experiments 
( Brodo,  1961b)  but  its  possible  importance  in  the  establishment  of  cer¬ 
tain  species  cannot  be  eliminated.  Since  virtually  no  work  has  been  done 
on  the  factors  involved  in  the  establishment  of  different  species  in  nature, 
little  can  be  said  about  this  important  aspect  of  lichen  ecology  at  this  time. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


37 


Certain  bark  characters  such  as  hardness  and  rate  of  exfoliation 
have  selective  effects  on  certain  lichens  and  certainly  cause  some  vertical 
lichen  zonation  (p.  22;  29-30).  Crustose  species  tend  to  be  most  abundant 
on  the  smooth,  young  bark  at  the  top  of  the  tree,  possibly  responding 
just  as  much  to  the  physical  bark  feature  itself  as  to  the  increased  light 
at  those  levels.  Hale  (1950,  1952a)  discusses  the  importance  of  bark 
texture  in  the  maintenance  of  certain  types  of  lichens  according  to  their 
anchorage  abilities.  Some  species  have  greater  abilities  to  reinvade  ex¬ 
foliating  bark  than  others  and  would  cause  vertical  zonation  along  a  bark- 
age  gradient.  One  then  might  view  a  single  tree  trunk  as  demonstrating  all 
stages  of  a  corticolous  succession  with  all  stages  in  time  frozen  at  differ¬ 
ent  levels  of  the  trunk. 

SUCCESSION 

Both  directional  and  nondirectional  changes  in  species  composition 
were  seen  within  certain  lichen  communities  (see  Hanson  and  Churchill, 
1961,  for  a  fairly  detailed  discussion  of  ecological  changes  of  different 
kinds).  The  tracing  method  used  in  the  growth  rate  studies  (see  Brodo, 
1965)  provided  a  means  for  demonstrating  the  fluctuations  in  local  lichen 
populations  and  the  constant  change  in  composition  and  coverage  of 
lichen  communities.  Reports  of  no  change  in  lichen  communities  in  up 
to  50  years  (A.  L.  Smith,  1921;  Cooper,  1928)  are  to  be  viewed  with 
some  skepticism  in  the  absence  of  precise  measurements  (as  pointed  out 
by  Smith,  1962).  Figure  16  presents  one  of  the  many  examples  of  fluctu¬ 
ations  which  were  observed.  Here  the  thalli  of  Parmelia  sulcata  are  shown 
to  grow  at  one  point,  while  in  other  places  they  fall  away  and  allow  the 
invasion  and  extension  of  Physcia  millegrana. 

When  populations  change  in  a  directional  fashion,  succession  can 
be  said  to  be  taking  place  (Hanson  and  Churchill,  1961).  The  changes 
described  below  may  be  truly  successional  or  may  be  the  first  stages  of 
a  cyclic  fluctuation.  These  changes  were  observed  in  the  growth  rate 
tracings  of  a  community  on  Quercus  alba  in  a  moderately  lighted  oak 
woods  (figure  17).  The  quadrat  was  at  a  height  of  one  meter  and  was 
facing  away  from  the  prevailing  wind  direction. 

1959:  Parmelia  sulcata  —  all  thalli  healthy,  robust. 

Physcia  millegrana  —  all  thalli  healthy  and  vigorous;  many 
very  small  thalli  present. 

Lecanora  caesiorubella  — -  one  thallus,  vigorous,  with  many 
large  apothecia. 

1961:  Parmelia  sulcata  —  some  thalli  showing  evidence  of  decay; 
most  healthy. 

Physcia  millegrana  —  all  thalli  healthy,  vigorous. 

Lecanora  caesiorubella  —  vigorous;  apothecia  unchanged. 

Dynamics: 

a.  Lecanora  caesiorubella  is  being  encroached  upon  and  cov¬ 
ered  on  all  sides  by  Physcia  millegrana,  although  both 
appear  to  be  healthy. 


38 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  16.  Population  changes  in  a  corticolous  lichen  community:  non- 
directional  shifts,  (a)  Parmelia  sulcata,  (b)  Physica  millegrana. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


39 


Figure  17.  Population  changes  in  a  corticolous  lichen  community:  direc¬ 
tional  shifts  (succession)  or  a  portion  of  a  cyclic  change,  (a)  Parmelia 
sulcata,  (b)  Physcia  millegrana,  (c)  Lecanora  caesioritbella  subsp. 
lathamii. 


40 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


b.  Wherever  Parmelia  sulcata  and  Physcia  millegrana  are 
both  healthy  and  are  growing  adjacent  to  one  another,  the 
Parmelia  is  growing  over  the  Physcia  with  one  exception 
in  a  very  local  area  of  a  Parmelia  sulcata  thallus. 

c.  Wherever  Parmelia  sulcata  appears  to  be  dying,  the  Physcia 
is  growing  over  the  Parmelia. 

d.  Small  regeneration  lobes  can  also  be  seen  in  the  dying 
areas  of  Parmelia  sulcata. 

1962:  Parmelia  sulcata  —  most  thalli  showing  considerable  decay. 

Physcia  millegrana  —  healthy,  vigorous. 

Lecanora  caesioruhella  —  half  the  thallus  whitened  and  de¬ 
caying. 

Dynamics: 

Physcia  millegrana  was  encroaching  considerably  on  the 
Lecanora. 

Succession  occurs  in  response  to  a  change  in  the  environment  of  the 
site  without  a  change  in  regional  climate  or  a  change  in  the  organism. 
The  rate  is  dependent  on  the  organism’s  rate  of  growth  and  the  environ¬ 
ment's  rate  of  change  ( Barkman,  1958).  The  sequence  of  the  successional 
stages  depends  on  the  characteristics  of  the  participating  organisms,  often 
both  physical  and  chemical.  Billings  and  Drew  (1938)  described  a  micro- 
succession  of  epiphytic  bryophytes  due  to  the  aging  of  the  bark  sub¬ 
strate.  Not  only  does  bark  change  in  time  due  to  the  tree’s  own  activity, 
but,  as  Barkman  (1958)  points  out,  the  epiphytes  themselves  alter  the 
moisture  capacity  and  acidity  of  the  bark.  The  forest  of  which  the  tree 
is  a  part  also  changes  in  time,  especially  with  reference  to  light  and 
humidity. 

With  both  corticolous  and  saxicolous  lichens,  the  successional  se¬ 
quence  is  usually  thought  of  as  crustose  to  foliose  to  fruticose  and/or 
bryophytes,  i.e.,  according  to  the  growth  form,  although  Rudolph  ( 1953a) 
has  pointed  out  many  exceptions  to  this  scheme.  The  sequence  described 
from  the  growth-rate  tracings  indeed  fits  into  the  more  standard  pattern. 
The  one  deviation  involves  poorly  developed  or  depauperate  specimens, 
in  which  case  the  succession  may  start  to  reverse.  Physcia  millegrana  is 
normally  overshadowed  by  Parmelia  sulcata,  except  when  the  latter  is  in 
poor  health,  at  which  time  the  Physcia  will  overgrow  the  Parmelia.  In 
areas  recently  subjected  to  air  pollution,  fruticose  and  foliose  lichens  are 
usually  more  severely  damaged  than  crustose  species,  and  succession  can 
therefore  be  reversed,  i.e.,  fruticose  to  foliose  to  leprose  species.  Bark¬ 
man  (1958)  described  such  a  reversal  in  Holland  with  Lobaria  giving 
way  to  mosses  which  finally  yield  to  Pleurococcus,  where  the  species 
were  found  near  industrial  centers.  This  is  thought  to  be  due  to  the  fact 
that  the  freer  an  organism  is  from  its  substrate  surface,  the  more  surface 
area  it  exposes  to  the  polluted  air  and  the  more  susceptible  it  is  to  air 
pollution.  Thus,  with  increasing  air  toxicity,  the  larger  foliose  and 
fruticose  lichens  will  be  the  first  to  go,  then  the  smaller  foliose  species. 


o 


island  localities  used  in  Brodo  (1961b)  are  numbers  1,  2,  3,  13, 
14.  The  north  shore,  red  oak  forest  localities  used  for  the  east-west 
transect  studies  are  numbers  1  through  12.  The  1961  north  shore 
transplant  experiment  was  set  up  in  localities  1,  2,  4,  10,  and  12. 


42 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


and  finally  the  leprose  crusts  and  algae.  In  New  York  City,  therefore,  the 
algal  communities  covering  the  trees  in  the  city  parks  can  be  considered 
to  be  in  a  disclimax  stage  maintained  by  air  pollution. 

Successional  stages  on  boulders  have  not  been  studied  on  Long  Island. 

On  the  inland  sand  habitats,  succession  often  proceeds  from  crusts 
such  as  Lecidea  uliginosa  or  L  granulosa,  to  Cladonia  spp.,  and  finally 
to  grasses  and  shrubs.  On  the  windswept  sand  dunes  of  the  south  shore, 
however,  the  reverse  is  often  observed.  Dune  grass  (Ammophila  breviligu- 
lata )  gains  a  foothold,  and  upon  its  death  and  decay  leaves  a  “stump” 
onto  which  some  species  of  Cladonia,  especially  C-  boryi,  can  become 
established.  The  clumps  of  Cladonia  will  close  over  the  sand,  more  or 
less  stabilizing  the  surface  and  will  decompose  and  provide  substrate  for 
other  plants  (Brodo,  1961a).  A  similar  but  more  nondirectional  cyclic 
change  was  described  by  Watt  (1947)  working  with  a  very  similar  com¬ 
munity:  Calluna  vulgaris,  Arctostaphylos  uva-ursi,  and  Cladonia  sylvatica 
( Cladonia  arbuscula).  In  his  scheme,  the  Cladonia  stage  can  give  way  to 
bare  soil,  again  upon  which  the  phanerograms  will  become  re-established. 
It  is  very  possible  that  the  same  cyclic  development  may  occur  on  the 
Long  Island  sand  dunes,  but  I  have  not  recorded  any  observations  to  that 
effect.  Alvin  (1960),  in  discussing  lichens  of  an  A m mcph ila  Call una 
dune  community  (p.  60),  considers  sand  stability  and  pH  as  prime  factors 
in  this  succession  with  reproductive  potential  (soredia  production)  as 
possibly  also  important. 

On  eroding,  sandy  loam  in  the  inland  portions  of  the  island,  a  type 
of  cyclic  change  can  be  seen.  Baeomyces  roseus  and/or  Pycnothelia 
papillaria  are  the  pioneers,  effectively  binding  the  soil  particles  together, 
providing  a  situation  suitable  for  the  invasion  of  many  other  species  of 
Cladonia  (particularly  O.  strepsilis,  C.  subcariosa,  and  C.  clavulifera) 
which  are  followed  by  grasses  and  herbs.  If  the  ground  is  disturbed  and 
subjected  to  new  erosion,  the  cycle  will  begin  again. 

I  studied  the  role  of  lichens  in  a  segment  of  old  field  succession 
over  a  period  of  3  years  at  the  American  Museum  of  Natural  History’s 
Kalbfleisch  Field  Research  Station  on  Long  Island.  Even  in  that  com¬ 
paratively  short  period,  some  interesting  trends  were  observed. 

The  field  under  study  (AP-5)  was  abandoned  in  1954,  and  was  thus 
6  years  old  when  these  observations  were  first  made.  In  1960,  the  phan¬ 
erogamic  vegetation  consisted  of  a  mixture  of  annua!  and  perennial 
weeds,  with  Andropogon  becoming  abundant  by  the  third  year  of  the 
observations.  In  general  the  trends  were: 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


43 


Vascular  Plants: 
Weeds  sparse; 
light  excellent 


Grass  becoming 
important; 
light  diffuse 


Y 

Grass  heavy; 
light  very  poor 


N on-vascular  Plants: 

1.  Pebbles  bare;  soil  exposed  around  weeds,  ex¬ 
cept  for  some  clumps  of  moss. 

2.  Pebbles  covered  with  pycnidia  of  Lecidea 
erratica,  some  non-podetiate  Cladonia  squam- 
ules  appear  on  moss  clumps  and,  to  a  lesser 
extent,  on  the  bare  soil. 

3.  Pebbles  covered  with  pycnidia  and  small  apo- 
thecia  of  Lecidea  erratica;  podetial  initials 
seen  on  Cladonia  thalli  in  moss  clumps  and, 
to  a  lesser  extent,  on  the  bare  soil. 

4.  Pebbles  covered  with  a  mixture  of  pycnidia 
and  mature  apothecia  of  Lecidia  erratica;  po- 
detia  of  Cladonia  cristatella  and  C.  chloro- 
phaea  well  developed  in  moss  clumps  and  on 
soil. 

5.  Rapid  decline  of  all  species,  especially  Cla¬ 
donia  spp. 

6.  Disappearance  of  Lecidea  erratica  from  peb¬ 
bles. 


Portions  of  this  succession  were  seen  in  various  quadrats,  and  the 
entire  successional  picture  is  actually  a  composite  of  the  many  segments 
observed. 

Robinson  (1959),  in  a  paper  dealing  with  old  field  succession  in 
North  Carolina,  also  noted  the  importance  of  Cladonia  cristatella  and 
C.  grayi  (C.  chlorophaea)  in  the  6-9  year  old  stage.  He  stated  that  the 
lichens  attain  their  greatest  dominance  after  the  decomposition  of  much 
of  the  moss  and  grass  vegetation.  If  the  same  sequence  follows  on  Long 
Island,  the  observations  at  the  Kalbfleisch  Station  could  represent  a  minor 
primary  succession  within  the  overall  successional  pattern  which  could 
only  be  seen  over  a  longer  period  of  time. 

Evans  and  Dahl  (1955)  noted  that  the  most  conspicuous  lichen 
cover  was  in  old  field  communities  of  mosses  and  perennial  weeds 
(“Bryoid  —  Antennaria  types”)  although  some  species,  including  Cla¬ 
donia  cristatella  and  C.  pyxidata,  attained  importance  in  the  “Poa  -  Aris- 
tida”  community.  The  Bryoid-Antennaria  community  is  well  lighted,  and 
becomes  established  on  dry,  unstable  soil;  whereas  the  Poa-Aristida  com¬ 
munity  is  slightly  more  shaded  and  is  found  on  more  stable  soil. 


44  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Because  of  the  common  occurrence  of  ground  fires  in  the  pine  areas 
of  central  Long  Island,  succession  on  burned  ground  and  bark  was 
studied  in  several  areas.  The  types  of  pioneers  on  burned-over  barrens 
depend  on  the  extent  of  the  fire.  If  even  very  limited  areas  are  left  un¬ 
burned,  a  large  number  of  species  may  be  available  for  reinvasion.  Fire 
can  get  very  close  to  a  lichen  colony  without  destroying  it.  In  southern 
New  Jersey  near  Tuckerton,  I  studied  an  area  which  was  burned  over 
not  more  than  2  years  previously.  The  fire  swept  through  the  area,  char¬ 
ring  almost  the  entire  ground  surface  as  well  as  many  tree  trunks.  The 
fire  apparently  was  windswept  and  very  rapid,  because  on  the  lee  sides 
of  many  trees  charred  bark  extended  to  a  height  of  about  4  feet,  whereas 
on  the  windward  sides  of  the  same  trees,  many  lichens  appeared  un¬ 
harmed.  On  the  soil,  a  similar  situation  was  seen.  Tiny  areas  untouched 
by  the  fire  supported  healthy  colonies  of  several  Cladonia  species,  par¬ 
ticularly  Cladonia  uncialis  and  C.  subtenuis  although  the  fire  had  devas¬ 
tated  areas  only  a  few  feet  away.  Small  moss  clumps,  especially  of 
Leucobryum  glaucum  or  Polytrichum  spp.,  seemed  to  provide  protection 
for  small  lichen  thallus  fragments,  and  some  reinvasion  of  the  surround¬ 
ing  area  probably  originated  from  these  clumps. 

With  a  rich  source  of  nearby  species,  succession  seems  to  be  rather 
haphazard  with  regard  to  pioneers,  and  is  mainly  dependent  on  which 
lichens  have  the  best  means  of  dispersal.  Charred  ground  is  soon  covered 
by  dust,  then  wind-blown  soil  and  other  plants,  and  is  recolonized  soon 
after  the  fire  has  gone.  Charred  bark,  however,  remains  uncolonized  for  a 
long  period  except  by  certain  specialized  species. 

Succession  on  an  area  almost  totally  destroyed  by  fire  gives  a  better 
indication  of  a  natural  succession  because  invasion,  with  few  excep¬ 
tions,  must  occur  from  outside  and  the  species  “selection”  is  much 
greater.  I  studied  such  an  area  in  Yaphank,  Long  Island,  adjoining  the 
Suffolk  County  Firematic  Training  Center.  The  fire  had  totally  destroyed 
the  ground  cover  and  charred  the  ground  over  an  area  of  about  50  acres 
or  more.  The  trunks  of  pines  were  burned  to  a  height  of  10-12  feet  and 
the  oaks  were  charred  to  a  lower  height.  Two  similar  areas  were  studied 
and  yielded  similar  observations.  One  was  in  Centereach  and  the  other 
near  Selden.  Prior  to  the  last  burn,  all  the  areas  were  pine-oak  barrens  of 
approximately  the  same  age. 

In  all  the  areas,  I  noted  that  lichens  invaded  the  soil  before  they 
invaded  the  bark  of  burned  trees.  The  uncontested  pioneer  was  the 
ubiquitous  Cladonia  cristatella,  followed  closely  by  C.  bacillaris  and  C. 
chlorophaea.  All  three  species  are  extremely  common  on  the  island  and 
all  have  very  wide  substrate  tolerances.  All  three  species  are  found  on 
tree  bark  of  many  types,  soil  of  many  types,  and  even  stones  and  gravel  if 
they  are  present.  Cladonia  caespiticia  was  observed  as  an  associated 
pioneer  in  the  Yaphank  and  Centereach  areas  as  well  as  on  charred 
ground  in  two  other  incompletely  burned  areas.  Sterile  thalli  of  Lecidea 
uliginosa  covered  small  patches  of  sand  in  the  Selden  locality. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


45 


No  reinvasion  of  the  charred  oak  bark  was  seen,  but  the  burned 
pine  bark  supported  a  number  of  species  of  crustose  lichens.  Lecanora 
subintricata ,  a  very  minute,  athalline  crustose  species,  was  collected  on 
a  completely  charred  and  almost  destroyed  pine;  Lecidea  anthracophila 
was  on  moderately  charred  bark  near  the  edge  of  the  burn.  Other  species 
such  as  Lecidea  scalaris  and  the  foliose  Panneliopsis  placorodia  were  on 
unburned  bark  just  above  charred  material  and  probably  were  remains  of 
a  pre-existing  population  rather  than  a  reinvading  one.  However,  Lecidea 
scalaris,  and  sometimes  L.  anthracophila,  have  been  collected  on  charred 
bark  on  numerous  occasions  and  undoubtedly  can  reinvade  recently 
burned  over  forests. 

I  did  not  study  the  long  term  effects  on  the  terrestrial  lichen  flora 
after  frequent  burning.  Buell  and  Cantlon  (1953),  however,  observed 
an  increase  in  the  lichen  cover  with  burning  frequency  over  a  period  of 
years.  Johnsen  (1959)  reported  a  slight  increase  in  lichen  cover  with 
periodic  burning,  but  declined  to  make  a  firm  statement  pending  more 
complete  data.  Both  the  above  studies  were  made  in  pine  forests,  the 
first  in  the  pine-oak  region  of  New  Jersey,  and  the  second  in  a  pure 
stand  of  loblolly  pine  (Pinas  taeda)  in  the  North  Carolina  piedmont. 

SPECIES  COMPOSITION  WITHIN  HABITATS 

It  has  long  been  known  to  field  botanists  that  certain  plants  tend  to 
be  found  in  association  with  certain  other  plants.  It  soon  became  con¬ 
venient,  therefore,  to  refer  to  these  groups  of  species  collectively  as 
“communities”  or  “unions.”  With  the  growth  of  the  field  of  phytosoci¬ 
ology,  hundreds  of  plant  communities  were  examined,  analyzed,  and 
named.  It  is  my  opinion  that  the  use  of  Latin  epithets  in  naming  bio¬ 
logical  communities  of  any  kind  implies  an  intricate  predictable  organi¬ 
zation  which  does  not  exist.  The  principles  underlying  this  opinion  as 
they  apply  to  lichen  communities  are  listed  below. 

1.  Each  local  lichen  population  has  definite  ecological  requirements 
(i.e.,  a  specific  niche);  for  certain  species  these  requirements  are  narrow, 
and  for  others  they  are  broad. 

2.  Lichens  with  similar  gross  ecological  requirements  will  tend  to  be 
found  together  more  frequently  than  lichens  with  dissimilar  ecological 
requirements.  The  more  similar  the  gross  requirements,  the  more  fre¬ 
quently  the  occurrence  of  the  two  together.  Since,  according  to  the  “eco¬ 
logical  exclusion  principle,”  no  two  species  having  exactly  the  same  niche 
requirements  can  exist  together,  as  the  ecological  requirements  of  two 
lichen  populations  approach  identity,  a  higher  and  higher  degree  of  com¬ 
petition  will  develop  between  them.  One  species,  once  present  in  a  habitat, 
could  successfully  exclude  an  ecologically  similar  species  by:  (a)  ex¬ 
tremely  rapid  growth  (preempting  suitable  available  space),  or  (b) 
chemically  or  physically  altering  the  habitat  preventing  the  establishment 
of  the  ecologically  similar  species.  This,  of  course,  could  be  effective 
only  if  the  establishment  requirements  of  a  species  were  different  from 


46 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  19.  Bark-borer  (a)  Assembled,  ready  for  use;  (b)  with  blade 
removed  to  show  additional  features  of  the  steel  holder. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


47 


its  survival  requirements.  Barkman  (1958,  p.  197)  believes  that  the 
production  of  a  growth  inhibitor  by  Opcgrapha  dubia  and  O.  cinerea 
prevent  the  two  from  occurring  side  by  side,  although  they  seem  to  have 
the  same  habitat  requirements. 

3.  Any  particular  area  or  locality  has  a  limited  set  of  potential 
species  available  for  colonization  due  to  that  locality’s  particular  ecologi¬ 
cal  and  phytogeographic  position.  For  example,  a  white  cedar  bog  on 
Long  Island  does  not  have  all  the  lichens  in  the  world  available  for 
colonization  (excluding  the  slim  possibility  of  chance  long  distance  dis¬ 
persal).  Only  those  species  whose  distributions  include  Long  Island 
which  had  means  to  arrive  at  Long  Island  (as  via  coastal  plain  swamps), 
and  which  require  or  can  to'erate  a  high  moisture,  low  light,  cool  tem¬ 
perature  environment  such  as  is  found  in  these  bogs,  could  occur  there. 
Thus,  out  of  approximately  16,000  species  known  to  science,  we  are  left 
with  about  30  which  may  be  found  in  a  Long  Island  bog.  On  any  par¬ 
ticular  tree  in  the  bog,  such  as  a  Chamaecy paris,  the  list  is  cut  down 
even  more  (eliminating  lichens  which  must  grow  on  the  ground,  rotting 
wood,  and  vegetation  or  smooth  bark,  etc.)  leaving  on'y  a  dozen  poten¬ 
tials.  The  chances  are  very  high  that  some  or  all  of  these  potential  species 
will  be  on  white  cedars  in  that  bog,  their  diaspores  having  arrived  there 
and  distributed  themselves  in  a  relatively  random  fashion. 

4.  The  conditions  in  any  particular  habitat  are  not  static,  mainly 
due  to  aging  of  the  substrate  itself  and  to  local  changes  in  microclimatic 
conditions;  consequently  the  community  composition  in  these  habitats  is 
not  static.  Succession  does  not  always  occur  with  the  same  sequence  of 
species  or  at  the  same  rate.  This  often  results  in  the  establishment  of 
mosaic  communities  of  mixed  development,  making  any  phytosociological 
classification  extremely  difficult. 

5.  The  composition  of  lichen  communities  varies  from  one  habitat 
to  another  in  an  unbroken  continuum  along  physical,  chemical,  or 
microclimatic  gradients.  Almborn  (1948)  cites  an  excellent  example  of 
such  a  continuum  following  an  illumination  gradient.  The  lichens  in¬ 
volved  were  members  of  a  community  on  Fagus. 

The  concept  of  “community”  as  used  here  should  not  be  confused 
with  the  integrated  biological  system,  consisting  of  lichens,  bryophytes, 
microorganisms,  vascular  plants,  and  animals,  of  which  it  is  a  part. 
Strictly  speaking,  one  can  even  think  of  each  lichen  thallus  as  a  sort  of 
“community”  ...  an  intimate,  highly  integrated  association  of  algae 
and  a  fungus. 

In  conclusion  then,  we  can  consider  a  “lichen  community”  to  be  a 
group  of  species  having  similar  gross  eco'ogical  requirements  and 
occupying  a  certain  habitat  together.  This  group  of  species  is  subject  to 
directional  and  non-directional  change  with  time  resulting  in  a  compo¬ 
sitional  continuum  from  one  group  to  another. 

If  we  decide  that  lichen  communities  should  not  receive  Latin 
names,  the  problem  of  how  to  deal  with  communities  still  remains.  One 


48 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


can  “classify”  a  community  according  to  its  floristic  composition,  as  do 
most  phytosociologists,  and  merely  refrain  from  giving  it  a  specific 
name,  or  one  can  delimit  communities  according  to  their  ecological 
affinities  by  classifying  their  habitats.  The  latter  method  is  employed 
in  this  work. 

Each  method  has  its  advantages  and  disadvantages,  and  a  final 
choice  depends  mainly  on  the  use  of  the  ultimate  product.  Barkman 
(1958)  in  discussing  epiphytic  communities,  ably  outlines  the  advan¬ 
tages  of  using  floristic  criteria  for  vegetational  analysis.  He  states  that 
phorophytes  cannot  he  used  alone,  since  “1.  the  kind  of  host  tree  is 
not  of  direct  influence,  2.  its  significance  varies  from  one  region  to  an¬ 
other,  3.  other  ecological  factors  are  thus  ignored  and,  last  but  not  least, 
4.  any  logical  system  should  be  classified  upon  the  characters  of  the  object 
to  be  classified,  in  casu  upon  the  vegetation  itself. . . His  arguments  are 
well  taken,  and  phorophytes  alone  are  not  used  in  the  system  outlined 
below.  However,  in  my  opinion,  (1)  phorophytes,  as  well  as  other  lichen 
substrates,  often  seem  to  have  a  distinct  influence  on  their  lichen  vege¬ 
tation  as  is  evidenced  by  the  number  of  species  which  are  wholly  or 
partially  substrate  specific  (p.  20-21),  (2)  the  changes  in  substrates  of 
certain  communities  from  one  area  to  another  often  give  important  clues 
pertaining  to  the  causes  of  accompanying  changes  in  vegetation,  (3)  eco¬ 
logical  factors  other  than  substrate  can  easily  be  included  if  needed,  and 
(4)  since  lichen  communities  themselves  are  basically  “unnatural,” 
i.e.,  they  are  only  fragments  of  true  biological  communities,  the  method 
of  community  classification  one  uses  should  depend  on  convenience  and 
usefulness.  The  chief  advantage  of  the  classification  of  habitats  over  the 
floristic  method  described  by  Barkman  is  that  the  former  does  not 
eliminate  any  vegetational  combination  and,  therefore,  makes  possible 
the  classification  of  a  total  flora  into  communities;  in  the  floristic  method, 
representative  associations  are  selected  from  the  total  flora,  leaving  many 
vegetational  combinations  not  considered. 

The  first  major  division  of  the  habitat  classification  which  follows  is 
by  the  various  vegetation  types  (in  their  broadest  concepts).  The  next 
division  is  by  substrate;  first  the  general  substrate  type  is  considered  and 
then  any  other  narrower  substrate  classification  that  seems  pertinent. 
Occasionally  a  microclimatic  division  is  made  beyond  that  of  the  substrate. 

Under  each  habitat  are  listed,  in  approximate  order  of  importance, 
lichens  which  have  a  high  probability  of  being  found  in  that  ecological 
situation.  These  species  comprise  the  “lichen  community.”  One  must 
keep  in  mind  that  the  species  lists  represent  potential  communities  and 
not  actual  ones.  Rarely  will  all  of  the  species  in  any  particular  community 
occur  together. 

Bark  characters  for  all  the  corticolous  community  phorophytes  are 
summarized  in  table  1. 

To  aid  the  reader  in  locating  specific  habitats,  an  outline  of  the 
habitat  types  precedes  the  discussions. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


49 


I.  Upland  Habitats 

A.  Corticolous 

1.  Pinus  rigida 

2.  Quercus  alba 

3.  Q.  prinus 

4.  Q.  velutina  group 

5.  Fagus  grandifolia 

6.  Acer  rubrum 

7.  Ulmits  americana 

B.  Saxicolous 

1.  Mortar  and  concrete 

2.  Granite  boulders 

3.  Pebbles  and  small  stones 

C.  Terricolous 

1.  Mossy  soil 

2.  Sandy  soil 

D.  Lignum 

1 .  Stable,  dry  lignum 

2.  Unstable,  highly 
decomposed  lignum 

II.  Bog  and  Swamp  Habitats 

A.  Corticolous 

1.  Chamaecyparis  thyoides 

2.  Acer  rubrum 

3.  Ilex  verticillata 

B.  Terricolous 

C.  Lignicolous 


III.  Maritime  Habitats 

A.  Aerohaline  stratum 
(salt  mist  zone) 

1.  Corticolous 

a.  Myrica  pensylvanica 
—  Primus  maritima 

b.  Primus  serotina 

c.  Juniperus  virginiana 

d.  Ilex  opaca 

2.  Saxicolous 

a.  Concrete  and  mortar 

b.  Granite  boulders 

3.  Terricolous 

a.  Stabilized  sand 

b.  Dune  sand 

4.  Lignicolous 

B.  Hygrohaline  stratum  (salt 
spray  and  storm  tide  zone) 

C.  Hydrohaline  stratum 
(littoral  zone) 


I.  UPLAND  HABITATS 
A.  Corticolous. 

1.  Pinus  rigida  (pitch  pine). 

Species:  (a)  base  —  Cladonia  bacillaris,  C.  incrassata,  C.  tris- 
tatella.  (b)  breast  height  —  Parmeliopsis placordia,  P.  aleurites, 
Lecidea  anthracophila,  L.  scalaris,  Bacidia  chlorococca. 
Comments:  The  best  lichen  development  occurs  on  the  edges  of 
the  bark  plates,  not  on  their  surface  (p.  28). 

Species  in  the  basal  community,  especially  Cladonia  incrassata  and 
C.  parasitica,  are  often  found  only  on  strongly  decayed  wood  and  on 
pine  bases.  The  reasons  may  lie  in  the  fact  that  both  substrates  are  very 
acid  (Barkman,  1958,  p.  113)  and  usually  moist.  Pine  needles  and 
bark  flakes  often  cluster  at  the  bases  of  pines,  forming  thick  piles  of 
material  which  retain  moisture  long  after  all  other  material  is  dry.  Thus, 
pine  bases  have  a  particularly  high  local  humidity. 

Pine  bark  species  found  at  breast  height  are  usually  narrowly  con¬ 
fined  to  pine  alone,  at  least  on  Long  Island,  although  pine-dwelling 


50  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

species  which  are  found  in  bogs  as  well  as  pine  forests  often  are  col¬ 
lected  on  Chamaecyparis  thyoides  or  even  Vaccinium  corymbosum. 

2.  Quercus  alba  (white  oak). 

Species:  (a)  base- — Parmelia  caperata,  Physcia  orbicularis , 
Cladonia  coniocraea.  (b)  breast  height  —  Parmelia  caperata, 
P.  rudecta,  P.  saxatilis,  Physcia  orbicularis,  Parmelia  sub- 
aurifera,  Physcia  millegrana. 

Comments:  The  relatively  high  moisture  capacity  and  low  acidity 
of  the  bark  of  Quercus  alba  renders  it  a  unique  habitat  in  the  black  oak 
and  pine-filled  forests  of  central  Long  Island  (see  also  Hale,  1955a). 
However,  its  lichen  vegetation  does  not  vary  much  from  that  of  black 
oaks,  with  a  few  important  exceptions,  notably  among  the  Physciae 
which  are  rather  common  on  white  oaks  and  rare  on  black  oaks.  Dis¬ 
tinctions  between  these  two  oaks  are  further  developed  under  the  discus¬ 
sion  of  the  black  oak  group.  LeBlanc’s  Parmelia  caperata,  P.  rudecta, 
P.  saxatilis,  Physcia  millegrana,  and  Ph.  orbicularis  unions  (LeBlanc, 
1963)  resemble  the  Long  Island  white  oak  bark  community  at  different 
points  in  the  continuum  of  lichen  composition. 

3.  Quercus  prinus  (chestnut  oak). 

Species:  (a)  base  —  Cladonia  coniocraea,  Parmelia  rudecta. 
(b)  breast  height  —  Parmelia  sulcata,  P.  rudecta,  P.  caperata. 

Comments:  The  very  hard,  impervious  bark  of  Quercus  prinus 
makes  it  a  rigorous  habitat  for  all  but  the  most  xeric  of  species,  especi¬ 
ally  above  the  base.  Its  relationship  with  the  lichen  vegetation  of  other 
trees  of  the  red  oak  forest  will  be  discussed  under  the  Quercus  velutina 
group. 

4.  Quercus  velutina  group  (black  oak  group)  including  Q.  velu¬ 
tina  (black  oak),  Q.  coccinea  (scarlet  oak),  Q.  rubra  (red 
oak),  and  all  hybrids,  especially  Q.  coccinea  X  rubra. 
Species:  (a)  base  — -  Cladonia  coniocraea,  C.  chlorophaea, 
Parmelia  caperata,  P.  rudecta,  P.  saxatilis.  (b)  breast  height 
(partial  shade)  - —  Parmelia  sulcata,  P.  rudecta,  Bacidia 
clilorococca,  Grapliis  scripta,  Lecanora  caesiorubella  subsp. 
lathamii,  Parmelia  caperata,  P.  saxatilis.  (c)  breast  height 
(light  good)  —  Parmelia  sulcata,  Lecanora  caesiorubella 
subsp.  lathamii,  Pertusaria  xanthodes,  Parmelia  subaurifera, 
Lecanora  chlarotera,  Parmelia  saxatilis,  Usnea  strigosa. 

Comments:  Comparisons  of  the  epiphytic  lichen  vegetation  of 
members  of  the  black  oak  group  indicate  that  all  species  support  very 
similar  communities.  Even  the  lichen  vegetation  on  Quercus  alba  bears 
many  resemblances  to  that  of  members  of  the  black  oak  group. 

Using  data  from  the  1961  transect  study  of  the  red  oak  forests  on 
the  north  shore  (p.  17),  the  epiphytic  lichen  communities  of  the  principal 
tree  species  were  compared.  Only  those  on  relatively  common  trees 
could  be  compared  statistically.  Kulcsinski’s  coefficient  of  community 
proved  to  be  the  most  useful  statistical  tool.  Only  stands  7  through  12 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


51 


(Sunken  Meadow  to  Shoreham )  were  used,  since  those  stands  west  of 
Sunken  Meadows  were  considered  under  the  influence  of  the  New  York 
City  atmospheric  conditions.  Where  it  seemed  valuable  and  pertinent, 
the  results  of  the  red  oak  forests  were  compared  with  those  of  the 
pine-oak  forests  derived  from  data  collected  in  1959  in  connection  with 
the  study  of  central  Long  Island  ( Brodo,  1961a).  Continuum  segments 
A  and  B  taken  together  are  considered  as  “pine-oak  forest”  for  the 
purposes  of  these  comparisons. 

It  is  possible  to  compare  the  epiphytic  lichen  floras  of  a  number  of 
trees  using  lichen  frequencies  at  1.3m  and  at  the  base,  as  did  Culberson 
(1955a),  or  by  disregarding  vertical  position.  The  latter  was  done  for 
the  lichen  communities  on  oaks  in  the  red  oak  forest.  The  communities 
were  then  arranged  in  sequence,  with  the  most  similar  closest  together 
and  those  most  dissimilar  farthest  apart.  Vertical  distribution  was  dis¬ 
regarded  in  this  case  since  there  were  few  differences  between  the  basal 
and  1.3m  vegetations  of  the  trees,  and  the  additional  lichen  species  intro¬ 
duced  by  the  combined  values  aided  in  the  computations.  The  matrix  of 
coefficient  values  with  the  ranked  communities  is  presented  in  table  4. 

The  most  striking  aspect  of  the  matrix  is  how  high  and  how  similar 
the  values  are.  In  Culberson’s  comparisons,  the  values  ranged  from  8  to 
76,  with  only  one  pair  of  tree  species  having  a  coefficient  over  70.  In 
the  Long  Island  study,  all  oaks,  particularly  Quercus  velutina,  Q.  coc- 
cinea,  and  Q.  coccinea  X  rubra  were  very  similar  in  their  epiphytic 
vegetation.  Only  Quercus  prinus  was  distinctly  apart  from  the  others. 

Table  4.  Degree  of  similarity  of  the  lichen  vegetation  growing  on  vari¬ 
ous  species  of  oak  in  the  red  oak  forest.  Coefficients  of  association  were 

based  on  the  formula  C=  — —  —  X  100,  where  a  =  the  number  of  lichen 

a  +  b 

species  on  one  tree,  b  =  the  number  of  lichen  species  on  the  compared 
tree,  and  w  =  the  number  of  species  found  in  common  on  both  trees. 
A  value  of  100  indicates  perfect  association  (i.e.,  identity,  as  far  as 
lichen  vegetation  is  concerned).  A  low  value  indicates  relatively  little 
similarity. 


Q.  vel. 

„  Q.  cocc. 

Q.  cocc.  14  , 

^  x  rubra 

Q.  alba 

Q.  rubra 

Q.  prinus 

Quercus 

velutina 

71  71 

68 

60 

42 

Q.  coccinea 

82 

73 

70 

46 

Q.  coccinea 
x  rubra 

14 

73 

46 

Q.  alba 

— 

— 

81 

61 

Q.  rubra 

— 

66 

Q.  prinus 

52 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Even  Quercus  alba  with  its  soft,  porous  bark,  was  almost  indistinguish¬ 
able  in  epiphytic  vegetation  from  the  black  oaks. 

These  results  agree  well  with  what  one  finds  in  the  actual  red  oak 
forest  —  a  rather  monotonous  and  sparse  epiphytic  flora  throughout 
the  stands,  regardless  of  the  phorophyte.  However,  in  the  pine-oak  forest, 
a  field  worker  is  struck  by  a  subtle  but  distinct  difference  between  the 
white  oak  and  black  oak  epiphytic  communities.  Analyzing  the  coeffi¬ 
cient  of  association  of  Q.  alba  and  Q.  velutina  communities  in  the  pine- 
oak  forest,  as  was  done  with  the  communities  on  oaks  in  the  red  oak 
forest,  we  arrive  at  a  figure  of  78,  which  indicates  they  are  similar  in 
their  epiphytic  flora.  Considering  the  basal  and  breast  height  lichen  fre¬ 
quencies  separately,  the  coefficient  of  association  values  are  74  and  75 
respectively,  still  not  reflecting  any  differences  between  the  trees.  Using 
only  the  presence  of  the  number  of  lichen  species  unweighted  by  the 
number  of  trees  examined,  rather  than  frequency,  one  arrives  at  a 
totally  different  and  much  more  realistic  picture  (table  5b). 

The  reader  may  come  to  suppose  that  this  is  merely  a  technique  of 
statistical  juggling  to  find  results  which  fit  preconceived  notions.  How¬ 
ever,  the  policy  of  looking  for  a  statistical  means  of  revealing  some  more 
or  less  apparent  ecological  phenomenon  actually  can  throw  a  great  deal 
of  light  on  the  real  factors  involved  in  this  phenomenon.  This  is  a  good 
case  in  point.  An  unweighted  species  presence  analysis  of  the  breast 
height  vegetation  of  Quercus  velutina  and  Q.  alba  reveals  that  the  two 
are  not  at  all  similar,  whereas  their  basal  communities  are  very  much  so- 
One  can  see  that  a  few  common  and  very  frequent  species  can  far  out¬ 
weigh  a  larger  number  of  rarer,  constant,  and  somewhat  substrate  specific 


Table  5.  Coefficients  of  association  of  lichen  vegetation  on  different 
tree  species  at  base  and  breast  height  quadrats.  The  coefficients  were 
computed  as  in  table  1. 

a.  RED  OAK  FOREST 


Base 

Breast  height 

Quercus  rubra  —  Q.  alba 

70 

67 

Q.  velutina  —  Q.  rubra 

81 

78 

Q.  velutina  —  Q.  alba 

83 

74 

b. 

PINE-OAK  FOREST 

Base 

Breast  height 

Q.  velutina  - —  Q.  alba 

83 

57(60)3 

:iSince  the  asymptote  of  the  species  sample  curve  for  the  Quercus  velutina  — 
breast  height  data  under  “pine-oak  forest”  was  not  as  sharp  as  was  seen 
with  the  other  curves,  an  extrapolation  from  45  to  ca.  90  trees  was  made, 
which  adds  approximately  2  species  to  the  Q.  velutina  flora.  It  can  be  assumed 
that  one  of  the  two  is  shared  with  Q.  alba  in  the  Q.  velutina-Q.  alba  compari¬ 
son.  raising  the  coefficient  from  57  to  60.  (See  full  discussion  in  text.) 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  53 

species.  For  example,  three  species  of  Physcia  and  two  of  Pertusaria  were 
found  only  on  Q.  alba  and  never  on  Q.  velutina. 

But  if  unweighted  species  presence  is  accepted  as  being  a  more 
instructive  test,  it  cannot  be  used  in  comparing  two  trees  of  different 
frequencies  unless  a  test  is  made  to  establish  the  extent  to  which  the 
sample  size  is  affecting  the  number  of  epiphytic  species  observed.  For 
example,  if  50  white  oaks  and  25  black  oaks  were  examined,  one  would 
normally  expect  many  more  epiphytic  species  on  the  former  if,  in  a  sam¬ 
ple  of  25  trees,  the  total  lichen  flora  on  that  tree  species  is  only  barely 
represented.  If  it  can  be  shown,  however,  that  in  this  example,  essen¬ 
tially  all  the  epiphytic  species  on  black  oak  were  examined  after  a 
sample  of  20  trees,  and  that  the  same  was  true  of  white  oak,  the  two 
trees  can  be  compared  without  regard  for  the  sample  size.  Species-sample 
curves  were  constructed  for  each  of  the  tree  species  and  their  lichen 
vegetation  (both  basal  and  breast  height)  in  the  red  oak  and  pine-oak 
forests,  and  it  was  shown  that  with  each  tree  species,  the  sample  size 
(40  to  90  trees)  was  sufficiently  large  to  allow  direct  comparisons  with 
other  trees. 

Coefficients  of  association  were  then  calculated  for  Quercus  velu- 
tina  and  Q.  alba ,  using  the  basal  vegetation  and  the  breast  height  vege¬ 
tation.  After  testing  sample  size  and  species  number  for  the  oaks  in  the 
red  oak  forests,  Q.  velutina  (including  Q.  coccinea.  as  in  the  pine-oak 
forest),  Q.  rubra,  and  Q.  alba  were  found  to  be  comparable  on  an  un¬ 
weighted  species-presence  basis  (table  5). 

Two  very  interesting  things  can  be  seen  from  the  table  of  coefficients. 
First,  the  breast  height  communities  of  Quercus  alba  and  Q.  velutina 
show  greater  difference  than  their  basal  communities.  Secondly,  this  dif¬ 
ference  appears  much  greater  in  the  pine-oak  forest  than  in  the  red 
oak  forest. 

Concerning  the  first  point,  it  should  be  noted  that  the  bark  of 
Quercus  alba  is  unlike  that  of  Q.  velutina  in  several  respects,  the  most 
obvious  being  hardness,  moisture  capacity,  and  color.  Bark  hardness 
was  not  measured  as  was  done  by  Culberson  (1955a),  but  white  oak  bark 
is  easily  flaked  off  and  gouged  with  a  fingernail,  and  the  bark  of  O.  velu¬ 
tina  is  sometimes  difficult  to  cut  into  even  with  a  sharpened  steel  knife. 
The  average  moisture  capacity  of  Q.  alba  was  found  to  be  69  percent 
(dry  weight),  41  percent  (volume),  and  14  percent  (area),  and  that 
of  Q.  velutina  was  found  to  be  38  percent  (dry  weight),  30  percent 
(volume)  and  17  percent  (area).  (See  p.  22-28  for  a  discussion  of  mois¬ 
ture  capacity  measurements.) 

Hale  (  1955a)  emphasized  the  differences  in  bark  characters,  par¬ 
ticularly  the  moisture  capacities,  between  Q.  alba  and  Q.  velutina.  Color 
is  more  important  than  would  at  first  be  suspected.  Heat  absorption  and 
thus,  indirectly,  evaporation  rate  must  be  influenced  by  bark  color.  All 
these  factors  add  up  to  a  characterization  of  black  oak  bark  as  a  very  dry 
habitat  (hard,  low  moisture  capacity,  high  evaporation  rate)  and  white 


54  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

oak  as  a  comparatively  moist  habitat  (soft,  high  moisture  capacity,  low 
evaporation  rate).  Many  authors  ( Barkman,  1958;  Billings  and  Drew, 
1938;  Young,  1938;  Brodo,  1959)  have  noted  the  great  importance  of 
the  physical  characteristics  of  bark  on  the  distribution  of  epiphytes.  This 
difference  in  moisture  relations  between  Q.  velutina  and  Q.  alba  would 
naturally  be  less  important  at  the  tree  base  where  the  microclimate  is 
normally  humid  (Barkman,  1958;  Billings  and  Drew,  1938)  than  at 
breast  height  where  microclimate  is  variable  and  usually  drier.  The  large 
number  of  species  on  Quercus  alba  may  thus  be  related  to  the  wetter 
microhabitat  which  can  support  a  greater  number  of  drought-sensitive 
species. 

This  difference  between  O.  velutina  and  Q.  alba  disappears  in  red 
oak  forests,  which  are  more  moist  and  more  shaded  than  pine-oak 
forests  (Brodo,  1961a),  and  where  the  vertical  zonation  is  in  general 
not  as  distinct  as  in  drier  habitats.  It  should  also  he  noted  that  many 
of  the  species  which  differentiated  the  two  lichen  communities  in  the 
we'l-hghted,  pine-oak  forest  were  photophilous  species  and  therefore  were 
absent  in  the  shaded  red  oak  forest. 

The  lichen  community  found  on  Carya  spp.,  not  included  in  the 
statistical  studies,  is  very  similar  to  that  of  the  younger,  smoother  parts 
of  the  oak  trunks,  especially  when  well-lighted  communities  are  compared. 

Depending  on  the  state  of  development  and  position  in  the  com¬ 
munity  continuum  (as  reflected  in  the  local  dominant  species),  the  fol¬ 
lowing  “unions”  of  LeBlanc  (1963)  can  be  referred  to  the  community 
found  on  black  oak  on  Long  Island:  Bacidia  chlorncocca  union,  Graphis 
scripta  union,  Parmelia  rudecta,  P.  saxatilis  and  P.  sulcata  unions. 

5.  Fagus  grandi folia  ( American  beech) . 

Species:  Trypethelium  virens,  Pyrenula  nitida,  Buellia  citr- 
tisii,  Graphis  scripta,  Phaeographis  dendritica. 

Comments:  The  hard,  smooth  bark  of  Fagus  grandifolia  effec- 
tive'y  limits  the  lichen  community  growing  on  it  to  crustose  species,  and 
the  den  e.  shade-producing  canopy  restricts  the  community  even  further. 

6.  Acer  rubrum  (red  maple). 

Species:  Hypogynwia  physodes,  Parmelia  perforata,  Pertu- 
saria  trachythallina,  Parmelia  subaurifera,  Lecanora  chlaro- 
tera,  L.  caesiorubella  subsp.  lathamii. 

Comments:  The  community  on  maple  in  upland  habitats  closely 
resembles  that  of  other  smooth,  hard  barked  trees,  especially  Quercus 
coccinea.  Hypogymnia  physodes,  however,  is  more  abundant  on  maples 
than  on  oaks  in  oak  forests. 

7.  Ulmus  americana  (American  elm). 

Species:  Xanthoria  parietina,  X.  fallax,  Physcia  millegrana, 
Ph.  stellaris. 

Comments:  Elms,  especially  as  they  occur  along  roadsides,  have 
been  studied  a  great  deal  in  relation  to  their  lichen  flora.  The  neutral 
pH  of  the  bark,  no  doubt,  has  a  direct  or  indirect  effect  upon  the 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


55 


epiphytic  flora,  because  other  neutral  barked  trees  (e.g.,  Populus  deltoides, 
Robinia  pseudoacacia )  have  very  similar  floras.  In  the  Braun-Blanquet 
system,  this  community  would  be  included  in  the  Xanthorion  parietinae 
Alliance  (Barkman,  1958).  In  LeBlanc's  treatment  of  Canadian  epiphytic 
communities  (LeBlanc,  1963),  the  Xanthoria  falla.x  union  would  most 
closely  apply  here. 

B.  Saxicolous. 

1.  Mortar  and  concrete.  pH  7-10.8 

Species:  (a)  Full  sun  —  Caloplaca  feracissima,  Lecanora 
dispersa.  (b)  Partial  or  full  shade  —  Caloplaca  flavovirescens, 
Placynthium  nigrum. 

Comments:  Mortar  and  concrete,  being  highly  alkaline  and  cal¬ 
careous,  are  equivalent  to  limestone  in  their  general  characters  and  in 
their  lichen  vegetation.  There  is  no  natural  occurrence  of  limestone  on 
Long  Island. 

2.  Granite  boulders.  Acidic,  coarsely  crystaline,  very  hard. 
Species:  (a)  Full  sun  —  Rinodina  oreina,  Lecanora  cinerea, 
Parmelia  arseneana,  P.  conspersa.  P.  stenophy-lla,  Sarcogyne 
clavus.  (b)  Partial  or  full  shade  —  Lecidea  albocaerulescens, 
Lepraria  zonata,  Parmelia  caperata.  Lecanora  cinerea.  Buellia 
stigmaea. 

Comments:  There  is  some  species-overlap  on  the  granite  commu¬ 
nities  of  well-illuminated  and  poorly-illuminated  boulders,  but  a  few 
species  are  absolutely  restricted  to  one  or  the  other  (e.g.,  Rinodina 
oreina  in  the  former  and  Lecidea  albocaerulescens  in  the  latter  com¬ 
munity). 

3.  Pebbles  and  small  stones.  Usually  smooth,  but  not  always; 
high  in  quartz.  Found  in  fields,  on  roadbanks,  or  in  other 
open  areas. 

Species:  Lecidea  erratica.  L.  coarctata.  L.  cyrtidia,  Acaro- 
spora  fuscata.  Rhizocarpon  obscuration. 

Comments:  Why  these  species  develop  on  pebbles  and  not  on 
boulders  of  similar  hardness  and  chemistry  is  hard  to  determine.  It  is  pos¬ 
sible  that  the  high  mineral  supply  (derived  from  seepage  and  splashing 
from  surrounding  soil)  or  higher  humidity  (due  to  close  proximity  to 
ground)  is  involved.  The  lack  of  stability  of  small  stones  and  pebbles 
is  also  probably  a  factor  in  limiting  the  kinds  of  lichens  which  can 
survive  in  this  community  (p.  29). 

C.  Terricolous. 

1.  Mossy  soil.  Gravelly,  but  relatively  rich  in  organic  matter; 
in  oak  woods  of  various  development;  pH  not  measured. 
Species:  Cladonia  subtenuis ,  C.  caespiticia,  C.  cristatella,  C. 
bacillaris,  C.  pleurota,  C.  chlorophaea,  C.  furcata. 

Comments:  This  community  is  best  developed  in  forest  glades,  or 
on  moss-covered  abandoned  roads.  It  is  almost  always  at  least  partially 
shaded. 


56  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

2.  Sandy  soil.  Little  to  no  organic  matter;  pH  4. 1-4.6. 

Species:  (a)  Unstable,  eroded,  sandy  loam  (roadbanks,  fire 
breaks,  etc.;  subsoil  or  very  fine  sandy  loam)  —  Baeomyces 
roseus,  Pycnothelia  papillaria,  Cladonia  strepsilis.  (b)  More  or 
less  stable  but  bare  sandy  loam  (slightly  more  sandy  than  “2”) 
— Cladonia  strepsilis,  C.  sitbcariosa,  C.  clavulifera,  C.  atlan- 
tica,  C.  chlorophaea,  C.  pleurota.  (c)  Very  sandy  soil — Lecidea 
uliginosa,  L.  granulosa,  Cladonia  cristatella,  C.  macilenta,  C. 
atlantica,  C.  uncialis,  Cetraria  islandica.  (d)  Dune  sand  — 
shifting,  sometimes  grass-covered  —  Cladonia  cristatella,  C. 
boryi,  C.  uncialis,  C.  submitis,  C.  chlorophaea,  C.  furcata 
(especially  f.  racemosa),  Cetraria  islandica  subsp.  crispa. 

Comments:  The  sandy-soil  communities  as  outlined  above  are 
largely  arbitrary  units  derived  from  a  continuum  formed  along  a  soil 
gradient  from  comparatively  rich,  sandy  loam  to  almost  pure  quartz 
dune  sand.  Although  a  few  species  are  more  or  less  confined  to  one  unit 
(e.g.,  Baeomyces  roseus  and  Pycnothelia  papillaria),  most  terricolous 
species  can  be  found  throughout  most  of  the  continuum. 

D.  Lignum. 

1.  Unstable,  dry  lignum  (fences,  planks,  decorticate  logs,  de¬ 
corticate  branches  and  twigs). 

Species:  Lecidea  aeruginosa,  L.  botryosa,  Cladonia  cristatella, 
C.  bacillaris,  Lecidea  myriocarpoides,  Bacidia  chlorococca. 

Comments:  This  community  occurs  in  both  partially  shaded  and 
sunny  habitats.  A  very  frequent  member  of  this  community  is  the 
imperfect  fungus  Coniosporium  olivaceum  Link. 

As  logs  and  planks  become  heavily  decomposed,  the  community 
composition  changes,  giving  rise  to  the  community  listed  below. 

2.  Unstable,  highly  decomposed  lignum  (rotting  logs  and 
stumps). 

Species:  Cladonia  parasitica,  C.  incrassata,  C.  bacillaris, 
Micarea  prasina. 

Comments:  The  characters  of  high  acidity  and  moisture  capacity 
seen  in  rotting  wood  have  much  in  common  with  the  bases  of  Pinus 
rigida  and  the  similarities  in  community  composition  are  obvious  (p.  49). 
The  community  is  best  developed  in  swamps  and  bogs  where  wood 
decays  quickly,  but  it  also  occurs  in  oak  forests  on  heavily  decomposed 
stumps. 

II.  BOG  AND  SWAMP  COMMUNITIES 

A.  Corticolous. 

1.  Chamaecy paris  thyoides  (swamp  white  cedar)  —  Vaccinium 
corymbosum  (highbush  blueberry). 

Species:  Parmelia  hypotropa,  Parmeliopsis  ambigua,  P. 
aleurites,  Cetraria  viridis,  C.  ciliaris,  Hypogymnia  physodes, 
Usnea  trichodea,  U.  subfusca  sensu  Motyka. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


57 


Comments:  The  corticolous  lichen  communities  on  these  two 
woody  bog  plants  are  remarkably  similar,  especially  in  view  of  the  fact 
that  one  is  a  conifer.  Several  members  of  the  bog  lichen  community 
occur  exclusively  in  bogs  and  only  on  these  substrates  (e.g.,  Cetraria 
viridis,  Parmeliopsis  ambigua,  and  Usnea  trichodea) .  The  relationship 
between  the  bog  habitat  and  the  bog  lichen  flora  will  be  discussed  later. 

Several  lichens  commonly  found  on  pine  in  pine  forests  are  found 
on  white  cedar  in  bogs  (Lecidea  anthracophila,  Ochrolechia  parella,  Par¬ 
meliopsis  aleurites).  Cetraria  ciliaris,  normally  found  on  white  cedars,  was 
collected  from  Betula  populifolia  in  two  different  maple  swamps.  The 
close  similarity  of  Betula  bark  to  that  of  conifer  bark  was  noted  by 
Barkman  (1958),  Skye  (1958),  and  others. 

2.  Acer  rubrum  (red  maple). 

Species:  Lobaria  pulmonaria,  L.  quercizaus,  Parmelia  rudecta, 
P.  caperata,  Pertusaria  ainara,  Bacidia  chlorococca. 

Comments:  The  smooth,  hard  bark  of  Acer  rubrum  undoubtedly 
has  a  great  effect  on  its  epiphytic  lichen  vegetation.  The  maple  commu¬ 
nity  is  almost  totally  different  from  that  on  bog  trees  and  shrubs  with 
looser,  more  absorbent  bark. 

3.  Ilex  verticillata  (black  alder). 

Species:  Trypethelium  virens,  Graphis  scripta,  Lecanora 
caesiorubella  subsp.  lathamii,  Pertusaria  xanthodes. 

Comments:  The  affinities  of  this  community  to  that  of  Ilex  opaca 
have  already  been  mentioned.  The  presence  of  the  other  species  men¬ 
tioned  may  well  be  due  to  the  better  light  conditions  in  Ilex  verticillata 
thickets.  The  dense  shade  produced  by  the  canopy  of  Ilex  opaca  exclude 
all  but  the  most  shade  tolerant  of  species. 

B.  Terricolous  (acid  boggy  sand,  edges  of  bogs). 

Species:  Cladonia  calycantha,  C.  atlantica. 

Comments:  This  community  also  is  well  developed  on  dry,  acid 
sand,  especially  as  found  in  pine  barrens. 

C.  Lignicolous  (rotting  logs). 

Species:  Cladonia  parasitica,  C.  incrassata,  C.  didyma,  C.  vul- 

canica,  C.  santensis,  C.  beaumontii. 

Comments:  The  community  on  rotting  wood  in  bogs  is  basically 
identical  with  that  of  drier  forests  except  for  the  occasional  presence  of 
the  four  rare  species  mentioned  last. 

A  marked  geographic  difference  was  seen  in  the  community  compo¬ 
sition  in  disjunct  localities  of  southern  New  Jersey  and  Cape  Cod.  The 
dominant  species  in  the  New  Jersey  bog-lignum  community  was  Cladonia 
santensis,  which  covered  large  areas  of  dead  wood  and  cedar  stumps. 
Cladonia  vulcanica  was  not  collected  there  at  all.  On  Cape  Cod,  Cladonia 
vulcanica  was  clearly  dominant  and  C.  santensis  was  not  collected.  The 
lignum  community  in  Long  Island  bogs  showed  neither  C.  vulcanica  nor 


58 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


C.  santensis  as  dominants;  both  are,  in  fact,  very  rare  on  the  island. 
Instead  Cladonia  incrassata  and  C.  parasitica,  both  common  throughout 
the  northeastern  coastal  plain,  were  most  conspicuous. 

III.  MARITIME  HABITATS 

Because  maritime  communities  are  so  heavily  influenced  by  their 
proximity  to  salt  water,  it  is  useful  to  classify  them  on  the  basis  of  their 
salt  water  exposure.  Des  Abbayes  (1934)  presented  a  detailed  discussion 
of  the  zonation  at  the  shoreline.  Following  Du  Rietz  (1925a  in  des 
Abbayes,  1934)  he  recognized  three  major  divisions,  (1)  the  aerohaline 
stratum,  which  is  strictly  terrestrial,  receiving  salt  only  as  a  fine  mist 
suspended  in  the  air  and  which  is  never  wet  with  salt  water,  (2)  the 
hygrohaline  stratum,  which  receives  salt  water  directly  as  salt  spray,  by 
immersion  at  very  high  tides,  or  at  the  high  spring  tides,  and  (3)  the 
hydrohaline  stratum,  which  is  submerged  with  every  high  tide  regardless 
of  the  season. 

Boyce  (1954)  and  Oosting  and  Billings  (1942)  measured  the  salt 
spray  concentrations  at  various  distances  from  the  mean  tide.  The  latter 
authors  found  that  salt  spray  is  greatest  on  the  exposed  side  of  the  fore 
dune,  less  at  the  hind  dune  summits,  still  less  at  the  lee  side  of  the  fore 
dune,  and  least  at  the  lee  side  of  the  hind  dune.  Boyce  reported  salt  con¬ 
centrations  of  up  to  2.2  mg  salt  dm2/hr.  at  a  distance  of  270  m  from 
mean  tide,  with  a  wind  speed  of  1 1  m/sec.  His  data  show  that  salt 
concentrations  closely  depend  on  wind  speeds,  as  well  as  distance  from 
the  salt  source.  On  Long  Island,  wind  speeds  of  1 1  m/sec,  are  very 
common  (p.  8)  and  so  one  can  safely  say  that  the  aerohaline  zone  ex¬ 
tends  at  least  270  m  from  the  water,  and  probably  much  beyond. 

Des  Abbayes  subdivided  the  hygrohaline  stratum  into  three  “eche¬ 
lons”  based  on  the  presence  of  certain  indicator  species.  Since  none  of 
his  indicator  species,  except  for  Verrucaria  microspora,  is  present  on 
Long  Island,  only  the  three  major  strata  will  be  used  in  this  community 
classification. 

A.  Aerohaline  stratum  (salt  mist  zone). 

1 .  Corticolous. 

a.  Myrica  pensylvanica  (bayberry)  —  Primus  maritima 
(beach  plum). 

Species:  (a)  Exposed  to  full  wind  and  salt-mist  (fore 
dunes,  bluff  tops,  beaches):  (1)  base  —  Parmelia  sulcata, 
P.  hypotropa;  (2)  breast  height  —  Rinodina  milliaria, 
Lecidea  various,  Parmelia  hypotropa.  (b)  Protected  from 
full  wind  and  salt  mist  (lee  side  of  dune,  groves  of  trees): 
(1)  base  - — -  Parmelia  sulcata,  P.  livida;  (2)  breast  height 
—  Parmelia  hypotropa,  P.  perforata,  Lecidea  various, 
Ramalina  fastigiata,  Usnea  strigosa. 

Comments:  All  the  species  listed  are  photophilous  with  high 
drought  and  salt  resistance.  It  is  evident,  however,  that  wherever  moisture 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


59 


is  greatest  in  the  dunes  areas,  the  lichen  vegetation  is  most  luxuriant. 
This  type  of  community  is  best  seen  in  the  Montauk  Point  area  and 
behind  the  moving  dunes  at  Promised  Land.  The  lichen  communities 
listed  above  occur  on  many  shrubs  along  the  shores,  and  almost  un¬ 
changed  on  many  of  the  dune  and  beach  trees.  (See  discussions  of  black 
cherry  and  red  cedar  below.) 

b.  Primus  serotina  (wild  black  cherry). 

Species:  Parmelia  sulcata,  P.  subaurifera,  Buellia  curtisii, 
B.  stillingiana,  Pertusaria  xanthodes,  Lecidea  various, 
Usnea  strigosa. 

Comments:  This  community  has  many  similarities  with  the  shrub 
communities,  and  differs  chiefly  in  the  inclusion  of  several  additional 
photophilous  crusts. 

c.  Juniperus  virginiana  (red  cedar). 

Species  :Physcia  millegrana,  P.  orbicularis,  Ramalina  wil- 
leyi,  Parmelia  hypotropa. 

Comments:  This  community  is  surprisingly  “nitrophytic”  (cf.  p. 
30)  perhaps  from  the  neutralizing  effects  of  salt  mist  (see  Barkman, 
1958). 

The  absence  of  conspicuous  crustose  species  is  perhaps  due  to  the 
instability  of  the  substrate. 

d.  Ilex  opaca  (American  holly). 

Species:  Trypethelium  virens,  Phaeographis  dendritica. 

Comments:  This  community  is  very  similar  to  that  on  Fagus  grandi- 
folia  and  Ilex  verticillata,  which  are  the  only  other  substrates  for  Trype¬ 
thelium  virens.  The  most  striking  similarity  between  the  trees  is  that 
all  three  possess  a  very  thin,  hard  outer  bark  with  a  living  layer  just 
beneath.  This  factor  alone  could  not  be  the  decisive  one  in  determining 
the  distribution  of  Trypethelium,  however,  since  many  other  trees  and 
shrubs  have  this  character  also  (e.g.,  Acer  rubrum  and  Amelanchier 
intermedia) . 

2.  Saxicolous. 

a.  Concrete  and  mortar. 

Species:  Xanthoria  parietina,  Caloplaca  citrina. 

Comments:  Verrucaria  muralis,  V.  nigrescens  and  Rinodina  salitia 

occur  as  rare  members  of  the  community  having  only  been  found  at 
Orient  Point. 

Xanthoria  parietina  and  Caloplaca  citrina  are  common  aerohaline 
species,  although  both  are  also  widely  distributed  far  from  salt  water 
(p.  249  and  252). 

Lecanora  dispersa  and  Candelariella  aurella  are  also  found  in  the 
aerohaline  stratum  as  facultative  members  of  the  community.  The  former 
is  listed  by  des  Abbayes  (1934)  as  a  typical  member  of  the  aerohaline 
stage  community.  Alvin  (1961)  noted  Catillaria  chalybeia,  Rinodina 
demissa  (R.  salina),  Lecanora  dispersa,  Xanthoria  parietina  and  Candel¬ 
ariella  vitellina  (ecologically  eauivalent  to  C.  aurella ?)  as  comprising  a 


60  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

community  found  on  the  bricks  of  a  sea  wall  on  the  east  coast  of 
England.  This  community  is  remarkably  similar  to  the  one  on  Long 
Island  except  for  the  absence  of  Caloplaca. 

b.  Granite  boulders. 

Comments:  No  lichens  were  seen  which  were  at  all  confined  to 
the  aerohaline  granitic  rocks,  although  several  species  normally  found 
farther  inland  were  found  growing  in  the  salt  spray  zone.  Parmelia 
caperata  and  Acarospora  fuscata  are  conspicuous  species  in  this  cate¬ 
gory.  Parmelia  caperata  was  listed  by  des  Abbayes  (1934)  as  common 
in  the  aerohaline  stage. 

3.  Terricolous. 

a.  Stabilized  sand  (as  on  Orient  Point). 

Species:  Cladonia  pyxidata,  C.  strepsilis. 

Comments:  On  Long  Island,  Cladonia  pyxidata  has  only  been 
collected  on  stabilized  beach  sand  in  the  salt  spray  zone.  It  is  interesting, 
from  the  standpoint  of  the  possible  salt-preference  of  this  species,  that 
I  have  seen  it  growing  in  luxuriant  abundance  on  beach  sand  on  the 
shore  of  Lake  Erie  (Point  Pelee,  Ontario).  Cladonia  strepsilis  is  clearly 
a  facultative  member  of  the  aerohaline  community. 

b.  Dune  sand  (as  on  Fire  Island  and  at  Napeague  Beach 
figure  9). 

Species:  Cladonia  submitis,  C.  boryi,  C.  uncialis,  C.  cris- 
tatella,  Cetraria  islandica,  Cladonia  chlorophaea. 

Comments:  Since  the  community  on  dune  sand  extends  essentially 
unchanged  into  inland  localities,  salt  spray  can  be  eliminated  as  im¬ 
portant  in  defining  its  distribution.  It  is  possible,  in  fact,  that  heavy  salt 
spray  such  as  would  occur  on  an  exposed  fore  dune  may  inhibit  the 
community’s  development  (p.  203). 

A  description  of  a  coastal  sand  dune  community  is  presented  by 
Alvin  (1960)  in  a  study  of  lichen  ecology  of  England’s  south  coast  at 
Dorset.  He  characterizes  the  dune  lichen  vegetation  using  a  “cross- 
section”  of  a  dune  much  as  I  did  with  a  south  shore  Long  Island  dune 
(Brodo,  1961a).  Although  Alvin’s  dune  system  was  more  complicated 
(consisting  of  three  ridges),  his  community  is  very  similar  structurally 
and  even  floristically  to  that  of  Long  Island.  Unbranched  Cladoniae 
such  as  C.  coniocraea.  C.  chlorophaea ,  and  C.  macilenta  were  closest  to 
the  ocean  on  relatively  unstable  sand,  giving  way  to  the  shrubby  Cladinae 
(C.  sylvatica,  C.  impexa,  C.  tenuis),  C.  furcata,  C.  uncialis,  and  Corni- 
cularia  aculeata  farther  back  in  protected  depressions  behind  the  first 
main  ridge.  Finally  appearing  on  the  stable  second  ridge  were  Lecidea 
(Biatora)  uliginosa,  Cladonia  crispata,  and  C.  squamosa.  With  a  few  species 
replacements  such  as  Cetraria  islandica  subsp.  crispa  for  Cornicularia 
aculeata  which  in  eastern  America  is  much  more  northern,  Cladonia 
submitis  for  the  more  northern  and/or  European  Cladinae,  and  the  North 
American  endemic  C.  atlantica  and  C.  squamosa  (which  is  more  mesic  on 
Long  Island),  the  dune  community  is  essentially  unchanged  in  structure. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


61 


This  is  but  another  example  of  closely  related  species  in  different  geo¬ 
graphic  areas  occupying  similar  niches  in  similar  habitats  to  create 
remarkably  similar  communities. 

4.  Lignicolous  (windswept  stumps). 

Species:  Lecanora  laevis. 

Comments:  Species  occurring  in  the  windswept  areas  of  the  island 
(on  beaches  and  sand  dunes)  often  are  very  well  developed.  Lecanora 
laevis  is  a  good  example,  often  covering  old,  hard,  windswept  stumps, 
especially  at  Orient  Point. 

B.  Hygrohaline  stratum  (salt  spray  and  storm  tide  zone) 
(saxicolous). 

Species:  Bacidia  umbrina,  Acarospora  fascata. 

Comments:  The  species  listed  as  “characteristic”  of  the  hygro¬ 
haline  community  (which  is  almost  non-existent  on  Long  Island)  are 
actually  far  from  their  normal  habitats  (farther  inland)  and  seem  to  be 
displaying  more  of  a  tolerance  for  the  zone  than  a  preference  for  it. 

C.  Hydrohaline  stratum  (littoral  zone)  (saxicolous)  (figure  14). 
Species:  Verrucaria  microspora,  V.  silicicola. 

Comments:  The  members  of  the  hydrohaline  community  are  found 
in  no  other  habitats.  Degelius  (1940)  reported  V.  microspora  from  the 
upper  hydrohaline  in  Maine.  Verrucaria  erichsenii,  which  Degelius  found 
abundant  in  the  lower  hygrohaline  stratum  was  not  found  on  Long  Island. 


City  Effect 

No  discussion  of  the  lichen  vegetation  of  a  partially  urban  area 
would  be  complete  without  a  consideration  of  the  detrimental  effects  of 
city  climate  on  lichen  growth  and  diversity  —  the  “city-effect.”  The 
city  effect  phenomenon  has  been  so  well  documented  in  recent  years  that 
it  has  become  one  of  the  most  well-known  of  lichen  characteristics,  even 
to  persons  knowing  little  else  about  these  organisms.  I  studied  the  prob¬ 
lem  as  it  occurs  on  Long  Island  in  some  detail  and  published  the  results 
in  a  recent  paper  (Brodo,  1966).  It  is  necessary  here  only  to  point  out 
certain  aspects  peculiar  to  the  Long  Island  situation,  and  to  repeat  the 
general  conclusions. 

Nearly  the  entire  western  third  of  Long  Island  is  made  up  of  a 
portion  of  New  York  City  and  its  heavily-populated  suburbs.  It  is  not 
surprising,  therefore,  that  a  strong  city  effect  can  be  demonstrated  on 
the  lichen  flora.  Unbiased  sampling  of  the  lichens  of  oak  forests  along 
the  north  shore,  transplant  experiments  of  foliose  lichens  along  two 
east-west  transects,  and  analyses  of  the  distributions  of  many  species  all 
were  made  to  determine  the  extent  of  this  effect.  The  results  of  these 
three  approaches  gave  slightly  different  types  of  information  but  basically 
corroborated  each  other. 

The  red-oak  forest  samples  of  epiphytic  lichens  showed  that  there 
is  a  gradual  increase  in  number  of  species  and  number  of  individuals  as 
one  leaves  the  city  center,  and  that  different  species  "recover”  at  different 
rates.  It  also  showed  that  there  is  little  “vertical  shift”  in  the  tree  bole 
vegetation;  i.e.,  lichens  normally  found  at  breast  height  on  trees  in  central 
Long  Island  do  not  gradually  become  more  and  more  restricted  to  tree 
bases  closer  to  the  city,  a  phenomenon  reported  by  several  observers  of 
city  lichens  in  other  parts  of  the  world  (Jones,  1952;  Rydzak,  1958; 
Gilbert,  1965). 

The  results  of  controlled  transplant  experiments  pointed  out  that  the 
city  environment  is  powerful  enough  to  kill  certain  foliose  lichens  within 
a  few  months  of  exposure,  and  that  with  a  year  of  exposure,  the  effect 
can  be  demonstrated  as  far  as  40  miles  from  central  Brooklyn. 

Distribution  maps  of  numerous  species  again  indicate  that  the  city 
effect  is  felt  up  to  about  40  miles  from  Brooklyn,  and  that  different 
species  enter  the  improved  flora  at  different  points. 

There  have  been  a  great  many  words  written  and  many  speculations 
made  concerning  the  causes  of  the  city  effect.  Air  pollution  is  the  factor 
most  often  given  as  the  main  element,  but  city-induced  drought  is  also 
cited  quite  often  and  certainly  must  be  considered.  There  is  little  doubt 
that  lichens  are  very  sensitive  to  air  pollutants,  (SOo  is  the  material  most 
often  mentioned),  and  few  would  deny  that  the  peculiar  biology  of 
lichens  makes  them  particularly  sensitive  to  changes  in  microclimate.  Why 
some  recent  workers  have  attempted  to  attribute  the  entire  effect  to  one 


63 


64  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

or  the  other  of  the  two  elements  is  therefore  puzzling  (see  Rydzak,  1958 
and  Gilbert,  1965). 

The  results  of  the  studies  on  Long  Island  were  compared  with  those 
of  previous  studies  done  in  Europe,  and  a  hypothesis  was  developed 
which,  1  believe,  incorporates  the  two  elements  of  city  environment  into 
a  plausible  explanation  of  the  city  effect.  A  detailed  discussion  of  this 
hypothesis  has  been  published.  In  general,  however,  it  appears  that  both 
city-induced  drought  and  air  pollution  affect  the  lichen  flora  of  an  urban 
area.  Air  pollution  is  carried  to  a  much  greater  distance  and  is  the 
chief  factor  in  the  reduction  in  species  diversity  over  long  distances. 
Drought  is  felt  only  locally,  in  and  immediately  around  the  most  built-up 
portions  of  the  city.  On  Long  Island,  the  pollution  effect  is  so  strong 
that  almost  all  lichens  are  killed  well  beyond  the  limit  of  the  drought 
effect.  The  influence  of  drought  is  best  demonstrated  in  the  field  by 
changes  in  the  vertical  distribution  of  corticolous  species.  This,  then, 
explains  why  there  is  no  vertical  distribution  shift  on  Long  Island,  while 
in  other  areas,  where  pollution  is  not  as  great  and  the  drought  and  pollu¬ 
tion  affected  lichens  overlap,  a  vertical  shift  is  noted  close  to  the  cities. 


Floristic  Elements 


INTRODUCTION 

In  an  area  as  small  and  geologically  uniform  as  Long  Island,  histori¬ 
cal  factors  cannot  explain  local  distribution  patterns,  since  ample  time  has 
been  available  for  the  uniform  distribution  of  any  plants  which  arrived  on 
the  island  other  than  very  recently  introduced  adventives.  The  migration 
routes  by  which  these  plants  reached  Long  Island  are  of  considerable 
interest,  however,  and  it  is  worthwhile  to  examine  some  of  the  probable 
sequences  of  events  which  fashioned  the  lichen  flora  of  Long  Island  as 
we  see  it  today. 

In  attempting  a  floristic  analysis  of  the  lichen  flora  of  Long  Island, 
it  has  been  necessary  to  analyze  the  major  distribution  patterns  repre¬ 
sented  in  eastern  North  America,  and  to  view  these  patterns  not  only 
with  regard  to  North  American  distributions  in  general,  but  also  with 
certain  aspects  of  worldwide  distribution.  There  is  a  much  greater  need 
for  a  broad  geographic  perspective  in  dealing  with  distribution  of  lichens 
as  compared  with  flowering  plants  since  endemism  on  a  species  level  is 
much  more  common  in  the  latter  (Ahti,  1964).  Approximately  24  per¬ 
cent4  of  the  lichens  of  Long  Island  are  endemic  to  North  America  as 
compared  with  an  estimated  65  percent  of  the  vascular  flora.  (The  vas¬ 
cular  plant  statistics  were  derived  from  an  unbiased  sample  from  Smith 
and  Ogden’s  unpublished  preliminary  flora  of  Suffolk  County,  in  conjunc- 
ton  with  comments  on  endemism  in  Fernald  [1950].) 

Many  authors  have  contributed  to  our  understanding  of  the  floristic 
patterns  to  be  seen  in  eastern  North  America.  Good  (1964)  provides  a 
general  pattern  of  the  major  elements.  The  forest  regions  of  the  eastern 
deciduous  forest  as  described  and  mapped  by  Braun  (1950),  although  not 
delineated  by  floristic  criteria,  reveal  some  of  the  basic  floristic  features 
of  eastern  North  America,  particularly  the  strong  influence  of  the 
Appalachian  Mountains. 

There  have  been  few  general  treatments  of  lichen  distribution  in 
North  America.  Thomson  (1963)  in  his  monograph  of  Physcia,  discussed 
American  distribution  patterns  with  an  emphasis  on  extra-American 
relationships.  Although  his  eight  categories  have  limitations  for  the  kind 
of  floristic  analysis  I  would  like  to  attempt  here,  two  of  Thomson’s  cate¬ 
gories  are  used  in  only  slightly  modified  form.  The  phytogeographic  sys¬ 
tem  proposed  by  Hale  (1961a)  is  very  useful  and  many  of  his  categories 
are  retained  essentially  unaltered. 

THE  CLASSIFICATION  OF  ELEMENTS 

The  floristic  elements  have  been  broadly  classified  according  to  gen¬ 
eral  climate.  An  Arctic-Boreal,  Temperate,  and  Tropical  element  can  thus 
be  recognized.  The  elements  are  each  divided  into  two  or  more  “sub- 

4  Computed  from  a  sample  of  87  percent  of  the  total  lichen  flora. 

65 


66 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


elements,”  and,  in  one  case,  further  subdivided  into  geographical  units. 
The  limits  of  these  categories  are  presented  below  and  representatives  of 
each  in  the  Long  Island  lichen  flora  are  listed  in  table  6. 

Element  I:  Arctic-Boreal. 

The  Arctic-Boreal  element  is  that  element  which  has  no  climatic 
northern  boundary.  Since  tree  line  would  be  a  northern  boundary  for 
arctic  corticolous  species  but  not  for  arctic  terricolous  species,  substrate 
was  bypassed  as  a  limiting  criterion. 

We  can  recognize  two  subelements  within  the  Arctic-Boreal  element. 
The  Arctic-alpine  subelement  corresponds  closely  with  Thomson’s  (1963) 
“circumboreal  arctic-alpine”  category.  It  is  distinctly  arctic  in  character, 
extending  into  temperate  United  States  only  in  the  alpine  zones  of  some 
of  the  eastern  and  western  mountains.  With  this  circumscription,  it  is 
obvious  that  no  member  of  this  subelement  could  be  present  on  Long 
Island.  The  members  of  the  Arctic-Boreal  element  which  do  extend  into 
boreal  and  temperate  climates  are  grouped  together  as  the  Boreal-tem¬ 
perate  subelement  (figure  20).  These  species  are  generally  very  wide¬ 
spread  due  to  their  broad  climatic  tolerances  and  access  to  circumpolar 
migration  routes. 

Element  IE  Temperate. 

In  the  Temperate  element  are  included  all  species  with  relatively 
distinct  northern  and  southern  climatic  limits,  usually  close  to  the  north¬ 
ern  and  southern  boundaries  of  the  United  States. 

The  temperate  element  can  be  divided  into  six  subelements.  The  first 
is  more  or  less  intermediate  between  typically  arctic  and  temperate  distri¬ 
butions.  This  North  Temperate  subelement  is  not  considered  as  arctic 
due  to  its  relatively  clear  northern  boundaries,  but  shows  distinct  boreal 
tendencies  in  many  instances.  It  is  best  developed  in  northern  United 
States  and  southern  Canada,  although  it  often  extends  southward  to 
include  most  of  continental  United  States  (figure  21  ).  The  “circumboreal 
north  temperate”  category  of  Thomson  (1963)  corresponds  to  this 
subelement,  which  includes  many  of  the  more  widely  distributed  common 
species. 

Three  important  physiographic  features  of  temperate  eastern  North 
America  are  the  coastal  plain,  the  Appalachian  and  Ozark  mountain 
systems,  and  the  Mississippi  valley.  The  Appalachian  Mountains  form  the 
core  of  the  area  occupied  by  the  Appalachian  subelement.  Extensions  and 
slight  modifications  of  the  basic  Appalachian  distribution  permit  us  to 
recognize  a  number  of  “units"  within  this  subelement.  The  Appalachian 
unit  includes  only  species  whose  basic  distribution  is  along  the  NE-SW 
mountain  chain  alone  (figure  22).  Extensions  to  include  the  Ozark 
Mountains,  the  Great  Lakes  region,  and  the  southern  Rocky  Mountains 
define  the  Appalachian-Ozark,  Appalachian-Great  Lakes ,  and  Appa¬ 
lachian-Great  Lakes-Rocky  Mountains  units,  respectively  (figures  23-25). 

Species  confined  to  any  or  all  three  of  the  segments  of  the  coastal 
plain  (i.e..  Gulf,  southern  Atlantic  and  northern  Atlantic)  are  included 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  67 

in  the  Coastal  Plain  subelement  (figure  26).  This  subelement  often  shows 
an  extension  into  the  Mississippi  Valley. 

A  large  number  of  eastern  temperate  species  are  not  restricted  to 
the  Appalachian  or  coastal  plain  regions  but  are  found  throughout  the 
eastern  deciduous  forest  from  the  Mississippi  Valley  (or  even  farther 
west)  to  the  Appalachians  or  the  east  coast.  These  species  comprise  the 
East  Temperate  subelement  (figure  27).  There  sometimes  is  a  distinct 
northern  or  southern  concentration  within  the  subelement  (see  maps  of 
Parmelia  galbina  and  P.  livid  a  in  Culberson,  1961)  but  its  division  into 
two  units  is  not  warranted. 

Often,  there  is  a  narrowly  restricted  concentration  of  records  in  the 
northeastern  states,  and  it  is  difficult  to  decide  whether  the  species  belongs 
to  an  eastern  segment  of  the  North  Temperate  subelement,  a  northern 
segment  of  the  East  Temperate  subelement,  or  a  portion  of  an  Appa¬ 
lachian-Great  Lakes  distribution.  Any  or  all  of  these  may  be  involved, 
of  course,  and  there  is  no  value  in  recognizing  separate  categories. 
“Northeast  Temperate”  species  have  arbitrarily  been  listed  with  the  East 
Temperate  subelement. 

There  are  a  number  of  wide-ranging  species  which  are  apparently 
relics  of  ancient  and  worldwide  distributions,  and  which  now  are  re¬ 
stricted  in  their  distributions  by  their  narrow  climatic  tolerances  (see 
below).  These  species  are  grouped  together  into  the  Oceanic  subelement. 
They  are  generally  characterized  in  North  America  by  having  east  coast- 
west  coast  disjunct  distributions  (figure  28).  Other  species  with  oceanic 
tendencies  (e.g.,  Lobaria  quercizans,  Collema  subfurvum)  but  which 
have  well  defined  distributions  in  one  of  the  subelements  already  de¬ 
scribed  are  considered  only  with  the  latter.  In  table  6  they  are  desig¬ 
nated  with  asterisks. 

The  oceanic  type  of  distribution  (usually  considered  as  an  element 
in  its  own  right)  perhaps  has  been  studied  more  than  any  other,  especi¬ 
ally  in  Europe  (Degelius,  1935;  Mitchell,  1961;  Faegri,  1958).  Degelius 
(1941)  also  made  some  observations  on  oceanic  species  in  eastern 
North  America. 

The  Oceanic  subelement  is  characterized  by  its  occupation  of  areas 
with  high  atmospheric  humidity  (although  degrees  of  rainfall  may  differ 
from  one  place  to  another),  and  areas  where  temperature  fluctuations 
are  small  from  one  season  to  another  (i.e.,  having  mild  winters  and 
cool  summers).  Though  areas  of  this  type  are  generally  coastal,  they  need 
not  be.  A  definite  oceanic  flora  can  be  found  in  the  Smoky  Mountain 
region  of  Tennessee  and  North  Carolina  (Degelius,  1941).  In  the 
present  study  no  distinction  is  made  between  “eu-oceanic”  (strictly 
oceanic  in  distribution)  and  “suboceanic”  (basically  oceanic  with  a  some¬ 
what  broader  tolerance  of  other  climates)  as  was  done  by  Degelius 
(1935)  since  lichen  distributional  and  ecological  limits  are  still  relatively 
poorly  known  within  North  America  as  compared  with  Europe. 


68 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  20.  Arctic-Boreal  element:  Boreal-Temperate  subelement.  Cla- 
donia  alpestris  (after  Ahti,  1961). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


69 


Figure  21.  Temperate  element;  North  Temperate  subelement.  Physcia 
stellaris  (after  Thomson,  1963). 


70 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  22.  Temperate  element;  Appalachian  subelement;  Appalachian 
unit.  P arinelia  appalachensis  (after  Culberson,  1962). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


71 


Figure  23.  Temperate  element;  Appalachian  subelement:  Appalachian- 
Ozark  unit.  Anzia  colpodes  (after  Hale,  1955c). 


72 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  24.  Temperate  element;  Appalachian  subelement;  Appalachian- 
Great  Lakes  unit.  Parmelia  olivetorum  (after  Culberson,  1958b). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


73 


Figure  25.  Temperate  element;  Appalachian  subelement;  Appalachian- 
Great  Lakes-Rocky  Mountain  unit.  Pseudevernia  furfuracea  (after 
Hale,  1955c). 


74 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  26.  Temperate  element;  Coastal  Plain  subelement.  Ramalina 
willeyi  (after  Howe,  1914). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


75 


Figure  27.  Temperate  element;  East  Temperate  subelement.  Parmelia 
aurulenta  (after  Hale,  1958). 


76 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Figure  28.  Temperate  element;  Oceanic  subelement.  Nephroma  laev 
gat um  (after  Wetmore,  1960). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


77 


Members  of  the  Maritime  subelement  are  restricted  by  habitat  avail¬ 
ability  to  the  temperate  maritime  zones  along  the  coast.  Compared  to  the 
maritime  flora  of  Europe,  this  subelement  is  very  poorly  developed  in 
eastern  North  America  (see  Degelius,  1940).  Although  the  Maritime 
subelement  theoretically  includes  species  restricted  to  the  sea  coast 
by  the  presence  of  salt  water,  salt  spray,  or  some  associated  marine 
influence,  no  aerohaline  species  from  Long  Island  appear  to  fit  into 
this  category. 

Element  III:  Tropical. 

Species  which  show  a  basically  tropical  distribution  are  grouped  into 
the  Tropical  element.  Representatives  in  the  western  hemisphere  are  usu¬ 
ally  widespread  in  Central  and/or  South  America  and  sometimes  can 
be  found  in  other  tropical  areas  throughout  the  world  as  well.  The  ele¬ 
ment  is  manifest  in  eastern  North  America,  centered  in  the  Appalachian 
mountain  system  and  on  the  coastal  plain,  and  is  thus  conveniently 
divided  into  Appalachian-Temperate  and  Coastal  Plain  subelements.  It  is 
perhaps  also  proper  to  recognize  an  Oceanic  subelement,  although  there 
appears  to  be  only  one  example  on  Long  Island. 

SUMMARY  OF  SIGNIFICANT  FEATURES 

Table  7  presents  a  summary  of  the  categorization  of  the  lichen  flora 
into  its  phytogeographic  elements  and  subelements,  with  figure  29  giving 
a  graphic  representation  of  some  of  the  important  facets  of  the  major 
categories.  The  summary  is  based  on  table  6,  which  includes  approxi¬ 
mately  81  percent  of  the  known  Long  Island  lichen  flora,  all  the  species 
for  which  we  have  some  good  phytogeographic  information. 

Some  observations  which  deserve  special  attention  are: 

1.  The  Arctic-Boreal  element  is  represented  by  21  percent  of  the 
flora,  all  but  two  species  being  partially  or  entirely  circumboreal. 

2.  Many  of  the  most  common  species  on  Long  Island  (see  table  7) 
are  members  of  the  Arctic-Boreal  element,  e.g.,  Cladonia  chlorophaea, 
C.  coniocraea,  Parmelia  sulcata  and  P.  saxatilis. 

3.  The  Temperate  element  is  most  abundantly  represented  (71  per¬ 
cent  of  the  flora). 

4.  All  North  American  endemic  species  are  in  the  Temperate  ele¬ 
ment,  mainly  in  the  East  Temperate  (6  percent)  Appalachian  (7  per¬ 
cent)  and  Coastal  Plain  (8  percent)  subelements.  In  all,  24  percent  of  the 
lichens  of  Long  Island  are  endemic. 

5.  Of  the  sampled  species  with  an  East  Asia-East  America  disjunct 
distribution  (16  in  all),  by  far  the  greatest  number  (38  percent)  are 
found  in  the  East  Temperate  subelement. 

6.  Considering  its  northern  latitude,  Long  Island  has  a  surprisingly 
good  representation  of  tropical  species  (8  percent).  Most  members  of  the 
Tropical  element  are  confined  to  the  coastal  plain  in  eastern  North 
America. 


PERCENT  OF  SPECIES  STUDIED 


78 


ARCTIC  -  TEMPERATE  TROPICAL 

BOREAL 


Figure  29.  Phytogeographic  affinities  of  the  Long  Island  lichen  flora. 
The  three  floristic  elements  are  depicted  on  the  abscissa.  Percentages 
on  the  ordinate  were  derived  from  a  sample  of  209  species  (81  per¬ 
cent  of  the  total  lichen  flora),  (a)  Species  also  found  in  Europe 
alone,  (b)  species  also  found  in  Asia  alone,  (c)  species  also  found 
in  both  Europe  and  Asia,  (d)  species  endemic  to  North  America. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


79 


7.  Most  of  the  species  having  amphiatlantic  distributions  (12  per¬ 
cent)  are  represented  in  the  Temperate  element  (the  East  Temperate  and 
North  Temperate  subelements). 


DISCUSSION 

Braun’s  (  1950)  map  of  the  forest  regions  and  sections  in  eastern 
North  America  reveals  that  Long  Island  lies  at  the  apex  of  three  major 
forest  types:  the  Oak-Chestnut  region,  with  its  origin  in  the  Appalachian 
foothills  and  southeastern  piedmont,  and  the  Southeastern  Evergreen 
forest  region,  which  lies  on  the  Gulf  and  Atlantic  coastal  plains.  The 
Hemlock-White  Pine-Northern  Hardwoods  region  lies  to  the  north  but 
is  separated  from  direct  continuity  with  Long  Island  by  an  area  of  Oak- 
Chestnut  forest  in  southern  Connecticut. 

The  new  vegetation  map  by  Kiichler  (1964)  shows  a  very  similar 
pattern,  but  with  the  vegetation  units  more  precisely  delimited.  For  ex¬ 
ample,  Kiichler’s  map  clearly  shows  the  change  from  the  oak-hickory-pine 
forest  of  the  southeastern  piedmont  to  the  northeastern  pine-oak  forest  of 
southern  New  Jersey,  Long  Island,  and  Cape  Cod,  and  shows  more  clearly 
the  differences  between  the  Gulf  coastal  plain  and  the  central-  and  north¬ 
eastern  coastal  plain  vegetation. 

Thus,  there  are  at  present  three  unbroken  biological  “highways” 
along  which  species  can  migrate  to  Long  Island  from  the  south,  and  an 
almost  uninterrupted  conifer-hardwoods  forest  to  the  north  which  pro¬ 
vides  easy  access  for  northern  species.  These  migration  routes  have 
existed  essentially  unchanged  for  thousands  of  years. 

The  greater  part  of  Long  Island  has  only  been  available  for  coloni¬ 
zation  since  late  Pleistocene  time  after  the  last  retreat  of  the  Wisconsin 
ice  in  that  area  (ca.  15,000  to  20,000  years  before  the  present).  It  is 
highly  probable  that  a  considerable  portion  of  the  northern  continental 
shelf  now  under  water  was  exposed  as  a  coastal  plain  during  and  just 
after  the  last  glacial  maximum  (Fogg,  1930;  Nichols,  1958).  As  the 
Long  Island  area  became  ice  free  this  extensive  coastal  plain  would  have 
provided  an  opportunity  for  unhindered  plant  immigrations  from  the 
south  and  west.  Soon  after,  sea  levels  rose,  due  to  the  melting  of  the 
glaciers  (Flint,  1957),  flooding  the  Long  Island  Sound  area  separating 
Long  Island  from  any  closer  connections  it  might  have  had  with  New 
Jersey  and  submerging  much  of  the  southeastern  New  England  coast. 
Much  of  the  submerged  coastline  north  of  Long  Island  reemerged  with 
the  up-doming  of  the  area  (Flint,  1957),  but  since  Long  Island  was 
apparently  on  or  just  south  of  the  “hinge  line”  it  remained  an  island. 

The  high  percentage  of  circumboreal  species  in  the  Boreal-Tem¬ 
perate  and  Northern  Temperate  subelements  (95  percent  and  77  percent, 
respectively)  is  not  surprising  in  view  of  the  extreme  likelihood  of  late 
Tertiary  and  Pleistocene  land  bridges  across  parts  of  the  arctic,  allow¬ 
ing  the  free  flow  of  plants  from  one  continent  to  another  (Fernald, 


80 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


1931;  Flint.  1957;  Colinvaux,  1964).  Graham  (1964)  cites  evidence 
for  North  Atlantic  migrations  via  the  arctic  islands  during  periods  of 
temperate  climate  in  the  Cenozoic.  Arctic  species  probably  can  still 
migrate  via  the  northern  islands  in  a  circumpolar  route  (Li,  1952). 

The  theory  of  pre-Pleistocene  continental  drift  is  still  very  much 
alive  and,  if  true,  may  explain  many  of  the  present  day  amphiatlantic 
lichen  distributions  ( Dansereau,  1957;  Good,  1964).  Hulten  (1958) 


Table  6.  Phytogeographic  categories  represented  in  the  Long  Island 
lichen  flora.  Aspects  of  each  species’  world-wide  distribution  are  noted 
as  follows:  A  =  found  in  Asia;  E  =  found  in  Europe;  N  =  North  Ameri¬ 
can  endemic;  X  =  not  endemic,  but  absent  from  Europe  and  Asia. 
Species  with  oceanic  tendencies  but  which  cannot  be  placed  in  the 
Oceanic  subelement  are  indicated  by  asterisks  (*).  Details  of  the  dis¬ 
tribution  of  each  species  and/or  references  to  published  summaries  or 
maps  are  presented  in  the  annotated  list. 

New  information  concerning  the  world-wide  distribution  of  a  few 
of  these  species  was  incorporated  into  the  following  list  too  late  to  be 
included  in  the  statistical  summaries.  There  is  no  appreciable  change 
in  any  of  the  summary  percentages. 


].  ARCTIC-BOREAL  ELEMENT 


A.  Arctic-alpine  subelement:  no  representatives  on  Long  Island 

B.  Boreal-temperate  subelement. 


Caloplaca  pyracea  AE 
Candetariella  aurella  AE 
C.  vitellina  AE 
Cetraria  islandica  AE 
Cladonia  alpestris  AE 
C.  arbuscula  AE 
C.  cariosa  AE 
C.  carneola  AE 

C.  chlorophaea  AE 
C.  deformis  AE 

C.  fmbriata  AE 

C.  furcata  AE 
C .  mitis  AE 
C.  pleurota  AE 

C.  pyxidata  AE 

C.  rangiferina  AE 
C.  scabriuscula  AE 
C.  squamosa  AE 
C.  uncialis  AE 
C.  verticillata  AE 
Dcrmatocarpon  miniatum  AE 
Diploschistes  scruposus  AE 


Hypogynmia  physodes  AE 
Lecanora  cinerea  AE 
L.  dispersa  AE 
L.  rubina  AE 
L.  symmicta  AE 
Lecidea  macrocarpa  AE 
Lecidea  vernalis  AE 
Parmelia  saxatilis  AE 
P.  sulcata  AE 
Parmeliopsis  ambigua  AE 
Peltigera  aphthosa  AE 
P.  canina  AE 
P.  polydactyla  AE 
P.  praetcxtata  AE 
Placynthium  nigrum  AE 

Rhizocarpon  grande  AE 

Rinodina  oreina  E 
Sarcogyne  simplex  AE 
Solorina  saccata  AE 
V errucaria  muralis  AE 

Xantboria  fallax  E 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


81 


II.  TEMPERATE  ELEMENT 


A.  North  Temperate  subelement 


Acarospora  fuscata  AE 

Lecidea  aeruginosa 

AE 

Alectoria  glabra  N 

L.  albocaerulescens 

AE 

Bacidia  umbrina  E 

L.  botryosa  AE 

Buellia  punctata  AE 

L.  coarc  tat  a  E 

B.  stillingiana  N 

L.  granulosa  AE 

B.  turgescens  N 

L.  nylanderi  AE 

Caloplaca  aurantiaca 

AE 

L.  scalaris  AE 

C.  cerina  AE 

L.  uliginosa  AE 

C.  citrina  AE 

L.  viridescens  AE 

C.  flavovirescens  AE 

Lobaria  pulmonaria * 

AE 

Candelaria  concolor 

AE 

Micarea  prasina  AE 

Catillaria  glauconigrans  N 

Parmelia  caperata  AE 

Cetraria  ciliaris  AE 

P.  conspersa  E 

Chaenotheca  phaeocephala  E 

P.  reticulata  AE 

Cladonia  bad  Haris  AE 

P.  stenophylla  AE 

Cladonia  coniocraea 

AE 

Pertusaria  amara  E 

C.  conista  AE 

Physcia  adscendens 

AE 

C.  macilenta  AE 

Ph.  aipolia  AE 

C.  multiformis  X 

Pli.  orbicularis  AE 

C.  nemoxyna  AE 

Ph.  s  tel  laris  AE 

Evernia  mesomorpha 

AE 

Rinodina  confragosa 

AE 

Graphis  scripta  AE 

R.  milliaria  N 

Lecanora  atra  AE 

Sarcogyne  clavus  E 

L.  hageni  AE 

Stereocaidon  saxatile 

E 

L.  muralis  AE 

Usnea  longissima  AE 

L.  varia  AE 

Verrucaria  nigrescens 

AE 

B.  East 

Temperate  subelement 

Bacidia  atrogrisea  AE 

Dimerella  diluta  AE 

B.  inundata  E 

D.  lutea  AE 

B.  schweinitzii  N 

Lecidea  anthracophila  E 

Buellia  curtisii  N 

L.  cyrtidia  N 

B.  polyspora  X 

L.  erratica  E 

Cladonia  apodocarpa  N 

Leptogium  corticola  E 

C.  brevis  E 

Leptogium  cyanescens  AE 

C.  caespiticia  AE 

Leptorhaphis  epidermidis  E 

C.  capitata  AE 

Micarea  melaena  AE 

C.  caroliniana  N 

Parmelia  aurulenta  A 

C.  clavulifera  A 

P.  galbina  A 

Cladonia  cristateUa  N 

P.  livida  N 

C.  floerkeana  AE 

P.  perforata  E 

C.  parasitica  AE 

P.  rudecta  A 

C.  strepsilis  AE 

Parmeliopsis  aleurites  AE 

C.  subcariosa  AE 

Phaeographis  dendritica  AE 

C.  subtenuis  X 

Physcia  millegrana  N 

82 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Physciu  subtilis  N 

Ph.  tribacoides  E 

Pycnothelia  papillaria  E 
Pyxine  sorediata  A 


Ramalina  fastigiata  AE 
Trypethelium  virens  N 
Usnea  mutabilis  N 
U.  strigosa  A 


C.  Appalachian  subelement 

1 .  A  ppalachian  unit: 

Buellia  dialyta  N  Haematomma  sp.  N 

B.  stigmaea  N  Parmelia  appalacliensis  N 

Cladonia  piedmontensis  N 

2.  Appalachian-Ozark  unit: 

Anzia  colpodes  X  Parmelia  hypotropa  AE 

Caloplaca  camptidia  N 


3.  Appalachian-Great  Lakes  unit: 


Alectoria  nidulifera  AE 
Anaptychia  palmulata  A 
Bacidia  chlorantha  N 
B.  chlorococca  E 
Baeomyces  roseus  AE 
Cetraria  tuckermanii  N 
Collema  subfurvum*  AE 


Haematomma  ochrophaeum  A 
Lobaria  quercizans *  N 
Parmelia  olivetorum  AE 
P.  subaurifera  AE 
Umbilicaria  mammulata  N 
U.  muhlenbergii  AE 


4.  Appalachian-Great  Lakes-Rocky  Mountain  unit: 


Cetrari  fendleri  N 
Cladonia  mateoeyatha  N 
Parmelia  subrudecta  AE 


Parmeliopsis  placorodia  N 
Pseudevernia  furfuracea  E 
Umbilicaria  papulosa  X 


D.  Coastal  Plain  subelement 


Bacidia  chlorosticta  N 

Lecanora  caesiorubella 

Cetraria  viridis  N 

subsp.  lalhamii  N 

Cladonia  atlantica  N 

L.  cupressi  N 

C.  beaumontii  N 

Melanotheca  cruenta 

N 

C.  boryi  A 

Parmelia  michauxiana 

N 

C.  evansii  N 

Pertusaria  propinqua 

N 

C.  floridana  N 

P.  xanthodes  N 

C.  incrassata  AE 

Porina  cestrensis  N 

C.  santensis  N 

Ramalina  stenospora 

N 

C.  simulata  N 

R.  willeyi  N 

C .  submitis  A 

Usnea  trichodea  A 

E. 

Oceanic  subelement 

Cladonia  terrae-novae 

N 5 

Xanthoria  parietina  AE 

Nephroma  laevigatum 

AE 

Xylographa  opegrapltella 

Pertusaria  velata  AE 

"This  species  was  added  to  the  Long  Island  list  too  late  to  be  included  in 
various  statistical  summaries  of  phytogeographic  affinities. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 
F.  Maritime  subelement 


83 


Verrucaria  microspora  E 


Verrucaria  silicicola  N 


III.  TROPICAL  ELEMENT 

A.  Coastal  Plain  subelement 


Cladortia  calycantha  AE 
C.  didvma  A 
C.  vulcanica  A 
Parmelia  dilatata  AE 


Pertusarici  tuberculifera  X 
Porina  nucula  E 
Ramalina  complanata  X 
T eloschistes  flavicans  AE 


B.  Appalachian-Temperate  subelement 


Anaptychia  obscurata  AE 
A .  pseudospeciosa  A 
Cladonia  cylindrica  A 
C.  pityrea  AE 


Pannaria  lurida  X 
Parmelia  perlata  AE 
P.  plittii  X 

Teloschistes  chrysoplithalmus  E 


C.  Oceanic  subelement 


Cladonia  carassensis  AE 


suggested  that  amphiatlantic  patterns  are  best  explained  by  postulating 
eastern  and  western  continental  migrations  from  the  Bering  Strait  region 
rather  than  trans-Atlantic  migrations.  Colinvaux  (1964)  has  sketched  the 
Pleistocene  floristic  activity  over  the  Bering  land  bridge.  Dahl  (1950) 
considered  present  day  American-European  disjunct  distributions  of 
lichens  and  some  other  plants  as  having  originated  from  arctic  parental 
populations  which  survived  the  ice  ages  in  unglaciated  areas  of  the 
arctic.  Among  the  Long  Island  lichens,  14  percent  of  the  species  have 
amphiatlantic  distributions. 

North  Temperate  and  some  Oceanic  species  possibly  migrated 
across  the  northern  regions  during  pre-,  inter-,  or  post-glacial  warm 
periods  and  later  retreated  southward  with  a  cooling  of  the  northern 
regions  and  the  glacial  advance,  returning  only  as  far  north  as  the 
northern  conifer-hardwoods  with  the  disappearance  of  the  ice.  Potzger 
(1952)  presented  palynological  evidence  to  suggest  that  the  pine  barrens 
of  southern  New  Jersey  served  as  a  refugium  for  many  boreal  commu¬ 
nities  which  were  displaced  southward  by  the  Wisconsin  glaciation. 
These  northern  plants  survived  the  ice  ages  side  by  side  with  southern 
communities,  only  to  migrate  northward  again  with  the  retreat  of  the 
ice.  Long  Island,  therefore,  was  in  an  excellent  position  to  be  invaded 
by  many  of  these  northern  species.  Cladonia  terrae-novae  probably  de¬ 
rived  its  distribution  pattern  in  this  way  (p.  201).  Possibly  some  North 
Temperate  species  also  were  introduced  from  Eurasia  into  the  North 
American  flora  during  the  post-glacial  warm  period  and  were  eliminated 
from  the  northern  boreal  and  arctic  latitudes  following  the  recent  cooling 
in  northern  climate. 


84 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Fernald  (1931)  and  Braun  (1955)  present  evidence  showing  that 
the  Appalachian  and  coastal  plain  floras  originated  from  pantropic  con¬ 
nections  that  invaded  the  Appalachians  at  a  very  early  time.  With  the 
uplift  of  the  area  during  the  Tertiary,  some  species  moved  out  onto  the 
newly  exposed  coastal  plain,  leaving  only  fragmentary  relics  behind  on 
the  Appalachian  plateaus.  Much  speciation  appears  to  have  occurred  in 
the  southern  Appalachians  during  the  long  period  of  its  isolation  (Fer¬ 
nald,  1931),  and  the  high  percentage  of  endemics  seen  in  the  Appa¬ 
lachian  and  Coastal  Plain  subelements  may  date  from  this  time. 

During  the  Pleistocene  glaciation,  coastal  plain  species  were  re¬ 
stricted  to  regions  south  of  the  ice,  although  probably  not  very  far  south 
(Braun,  1955;  Potzger,  1952).  With  the  retreat  of  the  ice,  the  northeast¬ 
ern  coastal  plain  became  available  for  colonization  from  the  south.  Fer¬ 
nald  (1931)  cited  much  botanical  evidence  to  support  his  theory  that 
there  was  a  post-glacial  period  of  relatively  warm  climate  when  the 
entire  coastal  plain  was  connected  by  a  continuous  land  formation  per¬ 
haps  as  far  north  as  Newfoundland  (see  also  Braun,  1955).  If  this  was 
the  case,  there  was  an  excellent  route  available  for  the  migration  of 
the  new  coastal  plain  species  northward  to  Long  Island  and  beyond  (cf. 
above,  and  Fogg,  1930). 

The  East  Temperate  subelement  had  at  least  two  origins:  one,  as  an 
eastern  segmentation  of  a  north  temperate  distribution,  and  the  other  as 
a  broadening  Appalachian  distribution.  Those  East  Temperate  species 
which  originated  from  the  north  are  likely  to  show  an  amphiatlantic 


Table  7.  Phytogeographic  affinities  of  Fong  Island  lichens.  All  per¬ 
centages  are  percent  of  total  sample  (209  species,  or  81  percent  of  total 
lichen  flora). 


Total  Percent 

of  of  In  Europe,  In  Asia,  Europe  N.  Araer. 
species  flora  not  Asia  not  Europe  and  Asia  Endemic 


# 

% 

if 

% 

# 

% 

# 

% 

I.  ARCTIC 

43 

21 

2 

1 

0 

0 

41 

20 

0 

0 

1.  Arctic-alpine 

2.  Boreal- 

0 

0 

0 

0 

0 

0 

0 

0 

0 

0 

temperate 

43 

21 

2 

1 

0 

0 

41 

20 

0 

0 

II.  TEMPERATE 

149 

71 

19 

9 

11 

5 

63 

30 

50 

24 

1.  N.  Temperate 

52 

25 

7 

3 

0 

0 

39 

19 

5 

2 

2.  E.  Temperate 

44 

21 

9 

4 

6 

3 

13 

6 

12 

6 

3.  Appalachian 

26 

12 

2 

1 

2 

1 

7 

3 

14 

7 

4.  Coastal  Plain 

2! 

10 

6 

0 

3 

1 

1 

1 

17 

8 

5.  Oceanic 

4 

2 

0 

0 

0 

0 

3 

1 

1 

1 

6.  Maritime 

2 

I 

i 

1 

0 

0 

0 

0 

1 

1 

III.  TROPICAL 

17 

8 

3 

1 

4 

2 

6 

3 

0 

0 

1.  Coastal  Plain 

8 

4 

2 

1 

2 

1 

2 

1 

0 

0 

2.  Appal. -temp. 

8 

4 

1 

1 

2 

1 

3 

1 

0 

0 

3.  Oceanic 

1 

1 

0 

0 

0 

0 

1 

1 

0 

0 

TOTALS: 

209 

100 

24 

11 

15 

7 

110 

53 

50 

24 

LICHENS  OF  LONG  ISLAND,  NEW  YORK 


85 


pattern,  whereas  those  coming  from  the  Appalachian  center  often  are 
either  North  American  endemics  or  show  evidence  of  a  widespread 
Tertiary  (and  East  Asia  disjunct)  distribution  (cf.  below). 

The  historic  relationships  of  the  various  elements,  suhelements,  and 
units  are  summarized  in  figure  30. 


< 

LU 


Figure  30.  Historic  relationships  between  floristic  elements,  subele¬ 
ments,  and  units  in  eastern  America.  Arrows  indicate  the  general 
direction  of  the  migration  of  species  from  one  area  (or  category)  to 
another.  The  categories  have  been  p'aced  in  quasi-geographical  posi¬ 
tions  relative  to  each  other.  Thickness  of  an  arrow  indicates  the 
relative  extent  of  the  migration;  a  dotted  line  arrow  refers  to  a 
slight  connection.  Tropical  and  Arctic-boreal  elements  indicate  their 
worldwide  affinities,  whereas  the  Temperate  element  is  relatively  iso¬ 
lated  except  through  its  tropical  or  boreal  connections. 


86 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


The  similarity  between  the  floras  of  temperate  eastern  North 
America  and  eastern  Asia  have  long  been  recognized  and  discussed  (Li, 
1952).  This  classical  disjunct  distribution  pattern  is  clearly  evident 
within  the  temperate  element  of  Long  Island  lichens.  Eight  percent  of 
the  Long  Island  flora  represents  Eastern  America-Eastern  Asia  disjuncts. 
Li  (1952)  states  that  Asian-Eastern  Temperate  floral  similarities  repre¬ 
sent  a  relic  distribution  of  a  Tertiary  flora  which  once  covered  the  tem¬ 
perate  to  arctic  northern  hemisphere.  The  fragmentation  of  the  flora 
was  caused  by  many  geological  changes  including  mountain  formations, 
continental  submergence,  climatic  change,  and  glaciation  (Li,  1952). 

It  is  especially  interesting  to  note  that  we  see  these  disjunct  patterns 
on  a  species  level  with  lichens,  whereas  phanerogamic  botanists  rely  on 
generic  similarities  (Fernald,  1931;  Li,  1952).  This  sort  of  evidence  can 
suggest  extreme  genetic  stability  and  slow  rate  of  evolution  in  many  lichen 
fungi  as  compared  with  flowering  plants  (see  also  Thomson,  1963).  In 
discussing  amphitropic  distributions,  Raven  (1963)  points  out  that  dis¬ 
junct  distributions  on  a  species  level,  especially  when  involving  autoga¬ 
mous  organisms  (as  would  be  the  case  with  lichens)  probably  are  due  to 
long  distance  dispersal,  particularly  by  migrating  birds,  and  not  to  any 
once  continuous  populations  which  became  extinct  in  intervening  areas. 
While  this  may  be  true  of  amphitropic  distributions  of  flowering  plants 
along  bird  migration  pathways,  it  is  hardly  possible  that  the  east  Asia 
disjunct  distributions  of  dozens  of  species  in  the  eastern  American  lichen 
flora  could  have  their  origin  by  long  distance  dispersal,  especially  when 
this  pattern  is  well  known  in  other  plants  at  higher  taxonomic  levels. 

Degelius’  wide  experience  with  the  European  lichen  flora  permitted 
him  to  recognize  a  number  of  European-American  vicariant  pairs  in  his 
studies  of  the  lichen  flora  of  Maine  (Degelius,  1940).  He  proposed  a  new 
category,  “subvicarious  species,”  to  include  species  which  do  not  entirely 
displace  each  other  but  instead  show  different  frequency  ratios  in  the 
different  areas.  He  suggested  various  alternate  possibilities  for  vicariant 
and  subvicariant  combinations  as  follows  (1-4).  Capital  letters  indicate 


the  species  is  abundant. 

and  small  letters 

indicate  it 

is  rare.  Alternatives 

5-11  have  been  added  and  will  be  discussed  below. 

A  Iternative 

N .  America 

Europe 

1. 

A 

B 

[  (true  vicariants) 

2. 

A  +  b 

B 

3. 

A 

a  +  B 

>  (subvicariants) 

4. 

A  +  b 

a+B 

5. 

a 

b  1 

(?) 

6. 

a 

B  \ 

7. 

A 

b  i 

1 

8. 

a  +  b 

B  ( 

(not  vicariants) 

9. 

A 

a  +  b  ; 

10. 

A  +  b 

A  ' 

| 

11. 

A 

A  +  b  1 

LICHENS  OF  LONG  ISLAND,  NEW  YORK 


87 


In  order  to  discuss  these  alternatives,  we  must  first  define  “vicariant 
(or  vicarious)  species.”  Vicariants  are  disjunct,  but  closely  related  spe¬ 
cies  which  are  similar  morphologically  and  often  ecologically.  I  think 
it  is  fair  to  say  that  most  definitions  implicitly  or  explicitly  assume 
approximately  equal  abundance  of  the  two  vicarious  populations.  This 
would  then  immediately  exclude  alternatives  6  through  11,  and  especi¬ 
ally  10  and  1 1  as  vicariants. 

Since  Degelius  almost  certainly  wanted  to  emphasize  relative 
abundance  rather  than  absolute  abundance  of  vicarious  pairs,  alterna¬ 
tive  5  is  superfluous  (being  equivalent  to  alternative  1)  and  can  be 
eliminated. 

There  are  many  other  possible  combinations  which  could  be  listed, 
of  course,  but  they  clearly  do  not  represent  vicariants. 

European-American  vicarious  species  found  in  the  Long  Island  flora 
are  listed  in  table  8.  Degelius’  use  of  Parmelia  {Pseudevernia)  cladonia 
and  P.  furfuracea  as  an  example  of  alternative  2  is  not  applicable. 
Pseudevernia  cladonia  is  relatively  rare  in  North  America,  while  P.  fur¬ 
furacea  is  more  widespread  and  often  common.  The  pair  would,  there¬ 
fore,  more  closely  fit  into  alternative  10  (assuming  the  North  American 
and  European  chemical  populations  of  P.  furfuracea  are  basically  con- 


Table  8.  European-American  vicarious  sub-generic  taxa  in  the  Long 
Island  lichen  flora.  In  the  cases  with  asterisks,  the  parent  or  daughter 
populations  have  apparently  continued  to  diverge  and  speciate,  producing 
double-taxon  vicariants.  The  problem,  while  slightly  more  complicated,  is 
basically  the  same.  Alternate  No.  1  of  Degelius  (1940)  refers  to  “true 
vicariants”  with  one  species  found  exclusively  in  America  and  the  other, 
equally  abundant,  found  only  in  Europe.  Alternative  No.  2  refers  to 
“sub-vicariants”,  with  the  European  species  represented  in  the  American 
flora  as  a  rare  or  very  local  plant  in  addition  to  the  more  abundant 
American  species. 


America 

Europe 

Degelius  ( 1940) 
Alternate  Number 

1.  Cladonia  subtenuis 

C .  tenuis 

2 

2.  C.  terrae-novae 

C.  impexa 

1 

3.  Pseudevernia  furfuracea 
(lecanoric  acid  strain) 

P.  furfuracea 
(olivetoric  and 
physodic  acid 
strains)* 

1 

4.  Lobaria  quercizans 

L.  amplissima 

2 

5.  Umbilicaria  papulosa 

U.  pustulata 

2 

88 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


specific)  and  should  no  longer  be  considered  as  vicariants. u  It  is  inter¬ 
esting  that  alternatives  3  and  4  are  entirely  absent.  Even  Degelius  (1940) 
could  not  give  an  example  of  No.  3,  and  his  example  of  No.  4  ( Lecanora 
carpinea-L.  pallida)  is  no  longer  applicable  in  the  light  of  recent  studies 
(Imshaug  and  Brodo,  1966).  A  consideration  of  the  origin  of  vicari- 
ants  and  their  probable  relative  abundance  is,  therefore,  of  interest. 

Vicariants  originate  from  speciation  of  an  isolated  portion  of  a 
widespread  population.  The  geographic  separation  of  a  parent  and 
daughter  species  may  occur  either  before  or  after  the  initiation  of  the 
new  species.  Love  (1955)  makes  a  definite  distinction  between  the  two 
types  of  resulting  vicariants.  He  regarded  those  species  having  arisen 
after  geographic  segmentation  of  a  parent  species  as  "true  vicariads,” 
and  those  species  which  arose  within  a  parent  population  by  some  imme¬ 
diate  genetic  isolating  mechanism  (such  as  polvploidization)  and  later 
became  separated  from  the  parent  population,  as  "false  vicariads.” 
Love,  after  discussing  the  usefulness  of  the  distinction,  explains  how,  in 
flowering  plants,  cytological  studies  can  establish  what  type  of  vicariism 
is  involved  in  each  particular  case.  Even  if  the  distinction  is  useful,  as  it 
may  well  be  in  certain  organisms,  i.e.,  in  lichen  fungi  which  appear  to  be 
genetically  "apomictic,”  the  distinction  cannot  be  made. 

In  any  case,  if  either  type  of  vicariism  mentioned  above  occurs,  it 
is  evident  that  alternatives  2  and  3  should  be  more  common  than  alterna¬ 
tive  1  because  of  the  low  probability  of  entirely  displacing  a  parent 
population  (i.e.,  with  the  parent  population  becoming  totally  extinct  in 
one  area).  It  is  therefore  significant  that  two  of  the  three  examples  of 
No.  1  cited  by  Degelius  (1940)  (Lobaria  quercizans-L.  amplissima; 
Umbilicaria  papitlosa-U.  pustulata)  now  appear  to  be  the  more  common 
alternative  2.  (It  should  be  pointed  out,  however,  that  the  North  Ameri¬ 
can  population  of  Lobaria  amplissima  is  disjunct  from  that  of  L ■  querci- 
zans,  being  known  only  from  southern  Mexico.) 

It  is  therefore  even  more  puzzling  that  there  are  no  examples  of 
alternative  3  in  the  lichen  flora.  One  could  hypothesize  that  all  lichen 
vicariants  are  “true”  vicariants  (sensu  Love,  1955)  and  have  come  from 
Europe  (suggesting  an  interesting  way  of  analyzing  a  migratory  direc¬ 
tion)  but  this  would  be  an  unlikely  conjecture  since  it  is  also  possible 
that  “false  vicariism”  is  involved  and  in  the  opposite  direction. 

Alternative  4,  which  requires  an  original  bidirectional  migration  or 
occasional  long  distance  imports  in  one  or  both  directions  with  the 
maintenance  of  an  equilibrium  ratio  between  the  two  species  (see  Mac- 
Arthur  and  Wilson,  1963)  appears  to  be  least  likely  of  all. 


'  Hale  (pers.  comm.)  informs  me  that  Pseudeveniia  cladonia  and  P.  furfuracea 
are  not  actually  closely  related,  and  that  the  former  is  locally  common  in 
some  areas  above  3000-4000  ft. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


89 


SUMMARY 

The  affinities  and  possible  origins  of  the  various  phytogeographic 
categories  are  presented  schematically  in  figure  30.  In  general,  there 
seems  to  have  been  two  routes  of  worldwide  distribution:  arctic-boreal 
and  tropical,  with  the  Temperate  element  largely  derived  from  one  of 
these  two  origins.  Tong  Island  is  approached  from  the  north  via  the  oak- 
chestnut  forests  which  included  parts  of  western  Tong  Island  before 
urbanization.  The  fragmentation  of  what  probably  once  were  continuous 
European-American  boreal  or  temperate  distributions  gave  rise  to  many 
examples  of  amphiatlantic  patterns,  including  several  vicarious  pairs  of 
species.  Many  northern  species  reached  the  island  from  the  south,  how¬ 
ever,  just  after  the  last  glacial  maximum.  Some  of  the  temperate  species 
which  originated  in  southeastern  United  States  reached  Long  Island  via 
the  Appalachian  Mountain  system,  which  partially  empties  out  into 
northern  New  lersey.  The  Atlantic  coastal  plain  provided  a  coastal 
“highway"’  along  which  southern  species,  many  of  which  originated  in  the 
southern  Appalachians,  could  migrate  northward  to  Long  Island.  These 
same  two  migration  routes  were  used  in  the  introduction  of  tropical 
species  into  the  Long  Island  flora.  Oceanic  species,  many  of  which  had 
ancient  origins  and  worldwide  distributions,  became  isolated  in  various 
areas  of  eastern  North  America  in  late  Tertiary  and  Quaternary  times, 
such  as  in  the  humid  and  comparatively  mild  Smoky  Mountains  of  the 
Appalachian  system  and  along  the  northeastern  coast  including  parts  of 
eastern  Long  Island,  Nantucket  Island,  Cape  Cod,  Newfoundland,  and 
Nova  Scotia. 


The  Lichen  Flora 


COLLECTIONS 

In  138  Long  Island  localities,  approximately  3,200  collections  were 
made.  An  additional  290  collections  in  southern  New  Jersey,  200  on 
Nantucket  Island,  and  400  on  Cape  Cod  provided  information  on  main¬ 
land  and  island  floristic  connections  with  the  Long  Island  lichen  vege¬ 
tation. 

Floristic  distribution  maps  often  come  under  serious  criticism  be¬ 
cause  they  are  said  to  represent  the  perambulations  of  the  collector 
rather  than  the  distribution  of  the  organisms.  To  overcome  this  short- 
occur.  As  a  result,  a  map  can  be  prepared  to  indicate  species  absence 
seen  in  a  locality  were  collected  no  matter  how  common  they  are.  In  this 
way,  a  determination  of  where  a  species  does  not  occur  can  be  made 
almost  as  accurately  as  the  determination  of  where  the  species  does 
occur.  As  a  result,  a  map  can  be  prepared  to  indicate  species  absence 
as  well  as  presence  (Imshaug,  1957a).  Of  course,  rare  species  will 
occasionally  be  missed  and  common  ones  will  occasionally  be  forgotten, 
but,  on  the  whole,  an  attempt  at  a  complete-collection  is  a  significant 
improvement  over  the  more  haphazard  collecting  methods  of  the  past. 
This  method  was  employed  in  all  the  Long  Island,  New  Jersey,  and  Cape 
Cod  localities. 

The  Long  Island  localities  are  listed  below  and  are  represented  by 
numbered  dots  in  figure  15.  In  the  interests  of  brevity,  localities  are  cited 
in  the  annotated  catalog  only  by  locality-number,  in  parentheses. 

KINGS  COUNTY:  (1)  Prospect  Park  (Brooklyn  Botanic  Gardens). 

QUEENS  COUNTY:  (2)  Forest  Park,  oak  woods;  (3)  Alley  Pond  Park, 
oak  woods  and  field. 

NASSAU  COUNTY:  (4)  Sands  Point,  shaded  maple-oak  woods  and  open 
field;  (5)  North  Hills,  dry  slope  above  swamp;  (6)  Valley  Stream,  Acer rubrum 
swamp  and  oak  clearing;  (7)  Rockville  Centre,  Hempstead  Lake  State  Park; 
(8)  East  Meadow,  "Hempstead  Plains;”  (9)  Brookville,  mature  oak  woods; 
(10)  Glen  Cove,  mature  red  oak-beech  woods;  (11)  Laurel  Hollow;  (12)  Cold 
Spring  Harbor,  path  and  black  oak  woods;  (13)  Cold  Spring  Harbor,  woods; 
(14)  Syosset-South  Huntington,  young  oak  woods;  (15)  Bethpage,  young  oak 
woods,  recently  burned;  (16)  Massapequa-Seaford,  black  oak  woods. 

SUFFOLK  COUNTY:  (17)  Centerport,  red  oak  -  chestnut  oak  woods 
and  roadside.  (18)  Vernon  Valley  (near  Northport),  red  oak  woods;  (19) 
South  Huntington-Half  Hollow,  oak-hickory  woods;  (20)  Dix  Hills,  oak  woods 
and  mossy  slope;  (21)  Commack,  mature  oak  woods;  (22)  Deer  Park,  oak 
woods  and  pine  woods;  (23)  Deer  Park,  woods,  swamp,  and  field;  (24)  near 
Babylon,  pine-oak  woods  bordering  acid  bog;  (25)  Captree  State  Park,  sand 
dunes;  (26)  near  King's  Park,  red  oak  woods;  (27)  San  Remo,  beech-oak-ash 
woods;  (28)  Hauppauge,  wet  woods;  (29)  Central  Islip,  young  oak  woods; 
(30)  Ronkonkoma,  oak-pine  woods;  (31)  Heckscher  State  Park  south  of  E. 
Islip,  oak-hickory  woods;  (32)  Oakdale,  young  oak  woods,  and  West  Sayville, 
roadside;  (33)  Fire  Island,  Cherry  Grove,  Sunken  Forest  Preserve,  Ilex  opaca 
grove:  (34)  Missequogue.  chestnut  oak-red  oak  woods;  (35)  St.  James,  red  oak- 


91 


92  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Figures  31-81.  Long  Island  distributions.  Each  open  circle  represents 
a  locality  where  a  collection  of  a  complete  set  of  lichen  species 
was  made,  but  where  the  species  in  question  was  absent.  With  the 
exception  of  figure  32  (see  below),  a  black  dot  indicates  that  a 
specimen  of  the  species  was  collected  in  that  locality.  Specimens 
collected  by  Latham,  Brainerd,  Hulst,  and  others  are  mapped 
whether  or  not  they  were  recollected  by  me  in  the  same  locality.  In 
some  cases  (e.g.,  the  pre  1900  New  York  City  collections),  these 
old  records  are  of  considerable  historic  interest.  All  Brooklyn  dots 
represent  pre  1900  collections. 

Figure  31.  Localities  of  oceanic  species.  Included  are  the  Long  Island 
localities  of  Cladonia  terrae-novae,  Collema  subfurvum,  Leptogium 
cyanescens,  Lobaria  pulmonaria,  L.  quercizans,  Nephroma  laevi- 
gatum,  ami  Pertusaria  velata.  Xanthoria  parietina  was  excluded 
since  its  distribution  appears  in  figure  84. 

Figure  32.  Bog  and  swamp  localities. 

Figures  33-41.  Lichens  found  mainly  in  bogs  and  swamps. 

Figures  42-52.  Lichens  found  mainly  in  pine-oak  forests.  (42-47:  with 
few  or  no  localities  east  of  Shinnecock;  48-52,  with  eastern  exten¬ 
sion;  42-45:  pine  specific;  46-47:  oak  specific.) 

Figures  53-56.  Lichens  found  mainly  in  morainal  areas.  (53-56:  terrico- 
lous;  57-60:  saxicolous;  61-63:  corticolous) 

Figures  64-70.  Lichens  found  mainly  in  the  humid  “fog  belt”  region. 
(69-70:  fog  belt  species  collected  in  New  York  City  prior  to  1870) 

Figure  71.  An  avoidance  of  the  red  oak  forest.  Physcia  millegrana. 

Figures  72-73.  The  scattered  distribution  of  two  terrestrial  lichens. 

Figures  74-76.  Lichens  found  mainly  on  sand  dunes  and  sand  plains. 

Figures  77-81.  Lichens  having  a  maritime  distribution.  (71-80:  aero- 
haline;  81:  hydrohaline.) 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


93 


94 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


95 


<r 


96 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


97 


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98 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


<r 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


99 


<r 


16  ?  MlLOM 


100 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


101 


<r 


102 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


61  *«  ?  KH.OM 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


103 


<r 


104 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


78  Rinodina  milliaria 


K)  MILES 
6i  I6  2KILOM 


O  o 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


105 


16  2  KILOM 


106 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


black  oak  woods;  (35  -  36)  Nesconset;  (36)  Centereach,  pine-oak  barrens 
burned  over;  (37)  Selden,  roadside;  (38)  Farmingville,  young  oak  woods; 
(39)  Patchogue,  young  oak  woods;  (40)  Patchogue,  open  pine  barren  recently 
burned;  (41)  Sayville,  wet  oak-pine  woods;  (42)  Old  Field,  dry  oak  woods; 
(43)  East  Setauket,  red-scarlet  oak  woods;  (44)  Port  Jefferson  Station,  dry  oak 
woods;  (45)  Coram,  mature  oak  woods;  (46)  Coram,  burned  over  pine  barren; 
(47)  Middle  Island,  young  oak-hickory  woods;  (48)  Patchogue,  field,  young 
oak  woods  and  maple  swamp;  (49)  Bellport,  open  field;  (S.  of  49)  Fire  Island 
opposite  Bellport,  between  dunes;  (50)  Miller  Place  -  Mount  Sinai,  red  oak- 
chestnut  oak  woods;  (S.  of  50)  Miller  Place,  oak-hickory  woods;  (51)  Middle 
Island,  mature  oak  woods;  (52)  Upton,  pine-oak  woods;  Ridge,  oak  and  pine 
woods;  (53)  Upton,  Brookhaven  National  Laboratory,  roadside  boulder,  oak 
woods;  (54)  Upton,  Brookhaven  National  Laboratory,  pine  woods  and  pine- 
oak  woods;  (55)  Yaphank.  oak  woods,  field  and  roadside  elm;  (56)  Brook¬ 
haven,  oak-pine  woods  and  bog;  (57)  Yaphank,  pine-scrub  oak  barren,  burned 
over;  (58)  Brookhaven  Station,  pine  barrens;  (59)  Shirley,  oak  woods;  (60) 
Fire  Island,  S.  of  Shirley,  sand  dunes;  (61)  Shoreham,  sand  bluffs  and  black 
oak  woods;  (62)  Shoreham-Wading  River,  shaded  oak-hickory  woods;  (63) 
Wading  River  Station,  old  black  oak  woods;  (64)  Montauk  Trail,  young  pine 
barren;  (65)  Upton,  young  black  oak  woods,  oak-pine  woods,  pine  woods  and 
maple  swamp;  (66)  Manorville,  open,  quaking  bog  and  surrounding  oak  woods; 
(67)  Manorville,  black  oak  woods  and  mature  oak  woods;  (68)  Manorville, 
oak  woods,  mature  oak  woods;  (69)  Manorville,  pine-oak  woods;  South 
Manor,  pine  barren;  (70)  South  Manor,  recently  burned  pine  barren  and 
young  oak  woods;  (71)  Center  Moriches,  black  oak  woods;  (NE.  of  71)  East- 
port,  graveyard;  (72)  Wading  River  (Wildwood  State  Park),  black  oak  woods 
and  bluffs;  (73)  near  Riverhead,  pine-oak  woods;  (74)  Calverton.  oak-pine 
woods  and  maple  swamp;  (75)  Calverton,  pine-oak  woods;  (76)  2  mi.  S.  of 
Calverton,  Bald  Hill,  pine  woods;  (77)  Riverhead,  black  oak  woods;  (78) 
Riverhead,  bogs  and  adjoining  oak  woods;  (79)  Riverhead,  pine  barren;  (80) 
Quogue-Riverhead  Rd.,  SW.  of  Flanders,  oak-pine  woods;  (81)  Riverhead. 
pine-oak  woods;  (82)  Eastport,  gravel  pit  bog;  (83)  Speonk,  pine-oak  barren, 
adjoining  maple  swamp  and  sphagnum  bog;  (84)  Remsenburg,  black  oak 
woods;  (85)  Riverhead,  pine  barren  and  young  pine-oak  woods;  (86)  Flanders, 
Chamaecyparis  bogs  and  pine  barrens;  (87)  Hampton  Bays,  pine-oak  woods 
and  Chamaecyparis  bog;  (88)  Quogue  Station,  oak  woods;  (89)  Quogue,  sand 
dunes;  (90A)  Northville,  deep  black  oak  woods;  (90B)  Mattituck,  sand  bluffs: 
(91)  Laurel,  oak-beech  woods;  (92)  South  Jamesport,  oak  woods;  (93)  Hamp¬ 
ton  Bays  (Squiretown ),  young  oak  woods,  and  Canoe  Place,  roadside  Caryci 
tomentosa;  (94)  Shinnecock  Hills;  (95)  Southampton,  Hudsonia- dune  area; 
(96)  near  Cutchogue  bluffs;  (97)  Peconic,  oak-hickory  woods;  (S.  of  97) 
Peconic  Station;  (98)  Southold  or  Laughing  Waters,  oak-hickory  woods; 
(99)  Noyack,  oak-hickory  woods;  (99-  111)  North  Haven;  (100  A)  Noyack, 
mature  oak  woods;  (100B)  Sag  Harbor,  oak-hickory  woods;  (101)  North 
Sea,  open  oak  woods;  (102)  North  Sea,  Chamaecyparis  bog  and  oak  woods 
above  bog;  (103)  Tuckahoe,  open  grassy  field;  (104)  Bridgehampton,  red 
maple  swamp;  (105)  Sagaponack,  sand  dune;  (106)  East  Marion,  oak-cherry- 
locust  woods  and  bluffs;  (107)  Shelter  Island,  Silver  Beach,  oak-hickory  woods; 
(SE.  of  107)  Shelter  Island,  Rt.  114  and  Smith  St.,  roadside;  (108)  Shelter 
Island,  Ram  Island  neck,  cherry-locust  woods;  (109)  Shelter  Island,  Ram 
Island  Drive,  red  cedar  thickets;  (110)  Shelter  Island,  Ram  Island,  oak-maple 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


107 


woods;  (111)  Shelter  Island,  N.  of  Nichols  Point,  open  oak  woods,  beech-oak 
woods,  beach  area;  (112)  Northwest,  oak-hickory  woods,  and  Three  Mile 
Harbor,  oak  woods,  open  fields  and  woody  bog;  (113)  Orient  Point,  red  cedar 
woods  and  shores;  (116)  Orient  Beach  State  Park;  (117)  Springs,  oak-hickory 
woods  and  oak  woods;  (118)  Springs,  roadside;  (119)  Amagansett,  oak-hick¬ 
ory  woods;  (120)  Napeague,  dunes  and  sand  flats  and  sand  barrens;  (121) 
Promised  Land,  sand  barrens,  oak  grove,  cherry  grove;  (122)  Gardiner’s 
Island,  field  and  old  oak  woods;  (123)  Gardiner’s  Island,  south  end,  grass¬ 
land;  (124)  Napeague,  sand  dunes  and  pine  barrens;  (125)  Hither  Hills  State 
Park,  pine  barrens  and  dunes;  (126)  Hither  Hills  State  Park,  exposed  ridge 
and  fresh  pond;  (127)  Hither  Hills  State  Park,  mature  oak  woods  and  wooded 
sand  bluffs;  (128)  Montauk,  white  oak-scarlet  oak  woods;  (129)  Montauk, 
grassy  downs;  (130)  Montauk,  low  sand  dunes;  (131)  Montauk  Point,  sand 
and  ridges;  (132)  Montauk  Point,  woods;  (133)  Montauk,  shaded  Ilex  ver- 
ticillata  thicket;  (134)  Fisher’s  Island. 

It  is  evident  from  the  map  in  figure  1 5  that  comparatively  few  col¬ 
lections  were  made  in  western  Long  Island.  In  Brooklyn,  Queens,  and 
Nassau  Counties,  collecting  areas  were  almost  exclusively  parks,  pre¬ 
serves,  or  highway  borders.  Even  in  many  parts  of  Suffolk  County,  par¬ 
ticularly  along  its  western  edge  and  along  the  north  shore,  the  only 
more  or  less  natural  areas  available  for  study  were  on  large  private 
estates  where  the  owners  were  kind  enough  to  allow  exploration  of 
their  property. 

ADDITIONAL  SPECIMENS  EXAMINED 

Several  herbaria  known  to  have  large  or  significant  Long  Island 
collections  were  visited,  including  the  Brooklyn  Botanic  Garden  (BKL) 
(Brainerd  and  Hulst  collections),  the  New  York  Botanical  Garden  (NY) 
(Torrey  Cladonia  collections),  the  Farlow  Herbarium  (FH)  (early 
Latham  collections  and  some  Ramalina  material  in  the  Howe  collections), 
the  New  York  State  Museum  (NYS)  (earliest  Latham  collections,  many 
reported  in  Burnham  and  Latham  [1914],  and  Charles  Peck  collections), 
the  Evans  herbarium  at  the  U.S.  National  Museum  (US:  Evans),  the 
University  of  Michigan  Herbarium  (MICH)  (Latham  collections  iden¬ 
tified  by  Fink),  the  Missouri  Botanical  Garden  (MO)  (Latham  collec¬ 
tions  identified  by  Dodge),  and  the  Herbarium  of  the  Staten  Island 
Institute  of  Arts  and  Science  (Staten  Island)  which  contains  several  old 
and  interesting  Long  Island  specimens.  A  few  specimens  were  also  seen 
from  the  University  of  Tennessee  herbarium  (TENN)  and  the  Cornell 
University  herbarium  (CUP).  Latham’s  personal  herbarium  is  given  the 
designation:  (Latham). 

TAXONOMY 

1.  Species  concept.  The  problem  of  “what  is  a  species,”  difficult  as 
it  is  with  any  group  of  organisms,  is  compounded  and  confounded  in 
lichens  by  the  fact  that  two  organisms  are  involved.  In  discussing  lichens, 


108 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


there  are  two  facets  to  the  problem:  (1)  what  do  we  mean  by  “lichen 
species”  —  the  consortium,  or  merely  the  lichen  fungal  component,  and 
(2)  the  common  problem  of  where  does  one  species  end  and  another 
begin. 

The  first  facet  was  solved,  in  theory,  in  1950  when  the  International 
Code  of  Botanical  Nomenclature  added  the  statement  “for  nomenclatural 
purposes  names  given  to  lichens  shall  be  considered  as  applying  to  their 
fungal  components”  (Lanjouw,  1961,  Art.  13,  Note  4).  Culberson 
(1961a)  seriously  challenged  that  position  and  maintained,  in  a  con¬ 
vincing  series  of  arguments,  that  the  name  of  a  lichen  should  apply  to  the 
entire  lichen  thallus  .  .  .  fungus  plus  alga.  His  main  arguments  center 
around  the  fact  that  almost  nothing  is  known  about  unlichenized  lichen 
fungi  and  that  the  little  that  is  known  points  to  the  fungi  as  being  quite 
different  in  morphology,  physiology,  and  ecology  from  the  lichen  as  a 
whole.  Since,  theoretically,  the  classification  and  identification  of  an 
organism  is  based  on  its  own  morphology,  etc.,  Culberson  asks  how  one 
can  apply  a  name  to  an  organism  based  on  the  totally  different  mor¬ 
phology,  physiology  and  chemistry  of  a  thallus  of  which  the  organism 
in  question  is  only  a  part. 

Although  his  arguments  are  well  taken,  I  still  believe  a  lichen  name 
should  refer  to  the  fungal  component  alone.  To  say  that  one  can  only 
classify  an  organism  divorced  from  all  other  members  of  its  biotic 
environment  is  not  valid.  Obligate  parasitic  fungi  are  studied  only  in  rela¬ 
tion  with  their  host,  and  yet  the  taxonomy  of  parasitic  fungi  has  not 
come  to  a  halt  because  of  it.  If,  perchance,  it  is  found  that  a  particular 
parasite  looks  different  or  has  different  reactions  on  different  hosts,  what 
may  be  thought  to  have  been  several  host  specific  species  at  one  time 
can  be  considered  to  be  one  species  later  with  no  particular  difficulty. 
Why  should  it  be  any  different  with  lichen  fungi?  I  believe  that  very 
few  different  lichens  will  be  found  to  have  the  same  lichen  fungus. 
Recently,  Uyenco  (1963)  showed  conclusively  that  the  morphology  of  the 
Coenogonium  lichen  thallus,  a  lichen  in  which  the  alga  is  the  dominant 
component,  is  due  to  the  fungal  component  alone.  She  showed  that  the 
same  lichen  fungus,  growing  symbiotically  with  different  species  of 
algae  in  different  regions,  will  produce  identical  lichen  thalli.  Thus, 
even  thallus  morphology  can  be  interpreted  as  a  fungus  character. 

To  say  that  lichen  chemistry  cannot  be  used  to  characterize  the 
fungal  component  of  a  lichen  is  to  disregard  the  genetic  basis  for  the 
ability  to  synthesize  a  lichen  acid.  The  lichen  fungus  is  involved  in  the 
production  of  the  chemical,  and  in  all  probability  at  most  derives 
certain  essential  chemical  precursors  from  the  alga  (Hess,  1959).  With 
a  growing  knowledge  of  the  biochemical  role  of  the  alga  in  a  lichen 
thallus,  we  will  probably  be  able  to  establish  a  system  in  which  the  un¬ 
lichenized  fungus  can  produce  characteristic  substances  in  culture.  Again 
we  see  that  a  thallus  character,  in  this  case  chemistry,  can  be  and  prob¬ 
ably  is  indicative  of  the  genotype  of  the  fungal  component. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


109 


It  therefore  seems  entirely  proper  to  use  thallus  characters  in 
characterizing  a  lichen  fungus.  It  also  seems  proper  to  use  the  name  of 
the  fungal  component  of  the  thallus  in  routine  references  to  the  thallus 
as  a  whole.  There  is  no  need  to  allow  lingual  gymnastics  to  confuse  and 
complicate  the  process  of  communication.  If  it  is  convenient  to  use  a 
fungus  name  to  refer  to  the  thallus  which  it  characterizes  in  nature,  so 
be  it.  All  those  involved  know  what  the  name  actually  stands  for,  and 
there  is  no  advantage  to  encumbering  discussions  with  constant  refer¬ 
ences  to  “Parmelia  sulcata  and  its  associated  algae,”  rather  than  just 
“Parmelia  sulcata ,”  the  lichenized  state  being  understood  unless  other¬ 
wise  specified. 

There  still  remains  the  problem  of  how  broad  or  narrow  a  species 
we  should  recognize  in  lichenology.  In  the  absence  of  evidence  for 
heterothallism  in  lichen  fungi,  objective  fertility  “tests”  as  applied  in 
phanerogamic  systematics  are  not  feasible,  and  so,  more  or  less  sub¬ 
jective  analysis  is  the  only  means  left  for  taxonomic  decisions.  It  has  been 
pointed  out  that  regarding  lichens  as  functional  “apomicts”  may  have 
some  merit,  especially  in  speciation  and  phytogeographic  considerations 
(John  Beaman,  pers.  comm.). 

I  think  it  is  fairly  obvious  that  generalizations  concerning  the  rela¬ 
tive  merits  of  specific  characters  cannot  be  made.  The  presence  of  soredia 
is  sometimes  important,  sometimes  unimportant;  certain  lichen  sub¬ 
stances  are  more  important  in  some  groups  and  less  important  in  others. 
This  problem  is  discussed  in  some  detail  by  Imshaug  and  Brodo  (1966) 
and  will  not  be  elaborated  on  here.  Suffice  to  say  that  the  more  informa¬ 
tion  we  have  about  a  species  and  its  close  relative,  i.e.,  the  distribution, 
morphology,  chemistry,  etc.,  the  easier  it  is  to  decide  whether  it  is  a 
species,  deserves  only  intraspecific  rank,  or  does  not  warrant  taxonomic 
recognition  at  all.  Thus,  for  Lecanora  caesiorubella,  the  rank  of  sub¬ 
species  was  selected  for  recognizable  segments  of  the  species  based  on  a 
great  deal  of  information  of  all  kinds.  With  less  complete  information, 
chemical  segregates  may  have  been  considered  “strains”  or  perhaps  full 
species.  The  recognition  of  some  species  here  is  tentative  pending  a 
more  extensive  and  intensive  investigation  of  their  group.  Such  species- 
pairs  as  Cladonia  didyma  -  C.  vulcanica,  and  C.  squamosa  -  C.  atlantica, 
the  C.  subcariosa  group,  and  others  need  more  work,  but  until  that  time 
the  narrow  limits  are  recognized. 

In  all  too  many  cases,  there  is  a  serious  question  as  to  the  status  of 
a  particular  taxon.  If  there  is  still  relatively  little  information  available 
on  which  to  base  a  firm  decision,  the  previous  treatment  which  I  con¬ 
sider  most  authoritative,  is  followed.  The  individual  systematic  problems 
of  various  taxa  are  discussed  in  detail  in  the  annotated  list. 

2.  Ecological  forms.  One  of  the  most  difficult  tasks  of  the  taxonomist 
is  to  determine  the  status  of  forms  found  in  differing  habitats  and  show¬ 
ing  different  morphological  or  chemical  characters.  For  example,  since 
both  moisture  and  light  are  needed  for  assimilation,  some  sort  of 


110 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


morphological  and  physiological  compromises  must  have  been  met  by 
the  lichens  in  their  adaptations  to  particular  niches.  But  has  the  change 
in  ecology  produced  the  changes  in  morphology,  or  does  the  morphologi¬ 
cal  variant  represent  a  genetically  stable  entity  confined  to  one  ecological 
habitat?  Weber  (1962)  recently  tried  to  answer  this  question  in  dealing 
with  the  ecological  modifications  of  some  crustose  lichens  in  the  south¬ 
western  United  States.  He  stressed  the  need  for  extensive  field  experi¬ 
ence  and  the  examination  of  large  numbers  of  specimens  in  making 
objective  decisions. 

In  some  cases,  the  situation  is  fairly  clear.  Xanthoria  parietina,  for 
example,  has  a  tendency  to  lose  (or  fail  to  develop?)  its  anthraquinone 
pigment  in  highly  shaded  places  (Thomson,  1949;  Barkman,  1958). 
Thalli  growing  on  concrete  blocks  at  Orient  Point  appeared  bright  yellow- 
orange  on  the  exposed  upper  surface  of  the  block  and  equally  vigorous 
but  a  pale  yellowish-white  on  the  shaded  side  of  the  block.  The  change 
in  habitat  from  strongly  insolated  to  shaded  (or  the  accompanying 
changes  of  dry  to  humid,  and  salt-sprayed  to  protected)  apparently  in¬ 
fluenced  the  quantitative  chemical  differences. 

Cladonia  cristatella  presents  a  somewhat  similar  situation.  When  in 
shaded  woods,  this  species  is  highly  branched  and  squamulose  with  a 
very  low  concentration  of  yellow  usnic  acid.  In  open  sunny  habitats,  the 
species  is  sparsely  branched,  almost  without  podetial  squamules,  and 
very  yellow  with  a  high  concentration  of  usnic  acid.  Increased  photo¬ 
synthetic  area  is  an  advantage  in  shaded  localities,  with  the  increased 
transpiration  from  the  increased  surface  area  being  insignificant  in  rela¬ 
tion  to  the  well-being  of  the  thallus.  In  exposed  areas,  since  light  is  not 
a  limiting  factor  and  moisture  is,  the  extra  surface  area  provided  by 
numerous  podetial  squamules  is  not  needed  and,  in  fact,  would  be 
disadvantageous  and  so  is  selected  against. 

The  production  of  extra  pigment  in  highly  illuminated  habitats 
applies  to  melanin  formation  as  well  as  usnic  acid  or  parietin  formation. 
Several  species  of  Cladonia,  particularly  C.  furcata  and  C.  atlantica,  show 
distinct  and  often  intense  browning  when  exposed  to  strong  sun.  Cetraria 
islandica  subsp.  crispa  shows  exactly  the  same  response  in  the  same  situ¬ 
ation.  Quispel  (1959)  and  Barkman  (1958)  suggest  that  lichen  pigments 
in  dry  thalli  may  have  a  role  in  the  protection  of  algae  from  high  light 
intensities  and  Rao  and  LeBlanc  (  1965)  presented  light  absorption  data 
supporting  this.  It  would  therefore  be  logical  to  expect  cortical  pigments 
to  be  in  higher  concentration  in  open  areas  than  in  shaded  areas. 

The  Peltigera  canina  group  provides  an  example  of  a  much  more 
difficult  problem.  There  is  basic  disagreement  on  the  status  of  ecologically 
differing  members  of  this  group,  particularly  P.  canina  sens.  str.  and 
P.  rufescens.  A  dry,  open,  eroded  habitat  is  characteristic  of  P.  rufescens 
whereas  a  more  cool,  moist,  mossy  habitat  is  typical  for  P.  canina. 
Thomson  (1950a)  maintains  that  there  are  all  gradations  from  one  type 
to  the  other,  and  that  P.  rufescens  is  merely  an  ecological  form. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  111 

Lindahl  (1953)  insists  that  the  two  are  clearly  separate  species.  He  per¬ 
formed  some  transplant  experiments  with  mature  plants  and  found  that 
the  transplanted  thalli  did  not  survive  well,  and  those  that  did  survive 
did  not  develop  into  the  type  characteristic  of  the  new  locality.  Lindahl's 
transplantations  were,  unfortunately,  not  controlled  with  thalli  trans¬ 
planted  to  similar  habitats  so  that  the  failure  of  his  plants  to  survive  in 
the  new  environment  is  not  significant  in  itself.  The  lack  of  morphologi¬ 
cal  change  in  a  mature  plant  is  also  to  be  expected  since  patterns  of 
growth,  once  having  reached  a  “point  of  no  return”  (Cantino,  1961), 
may  be  difficult  if  not  impossible  to  alter.  Transplantations  of  isidia 
or  tiny  squamules  may  prove  to  be  valuable  in  determining  the  role  of 
ecological  conditions  on  the  thallus  forms. 

3.  Infraspecific  laxa.  The  use  of  infraspecific  categories  in  this 
paper  is  admittedly  erratic.  In  general,  varieties  and  forms  are  not  con¬ 
sidered  and  no  new  infraspecific  taxa  are  described,  although  a  few  new 
combinations  are  made  involving  varieties.  The  few  exceptions  involve 
references  to  more  clearly  defined  taxa  which  sometimes  are  considered 
as  full  species  (such  as  Rhizocarpon  obscuration  f.  reduction )  or  thor¬ 
oughly  studied  taxa  which  fit  well  into  an  infraspecific  rank  (such  as 
Lecanora  caesiorubella  subsp.  lathamii) .  The  numerous  varieties  and 
forms  described  in  Cladonia  are  not  recognized,  since  the  large  majority 
are  undoubtedly  growth  forms  and  ecological  variants  and  the  rest  have 
been  insufficiently  studied. 

4.  Keys  and  annotated  list.  The  arrangement  of  the  flora  into  sub¬ 
classes  and  orders  follows  Hale  and  Culberson  (1966).  The  family  con¬ 
cepts  follow  Hale  (1961a)  with  the  following  exceptions:  the  Nephro- 
maceae  (after  Wetmore,  1960),  the  Baeomycetaceae  (after  Rasanen, 
1943),  the  Candelariaceae  (after  Hakulinen,  1954),  and  the  Teloschis- 
taceae  and  Physciaceae  (after  Nannfeldt,  1932).  The  arrangement  of 
genera  within  the  families  follows  Zahlbruckner  (1926b).  Species  have 
been  placed  in  alphabetical  order  within  the  genera  except  for  Cladonia, 
which  was  arranged  according  to  Mattick’s  (1940)  (and  in  the  case 
of  the  subgenus  Cladina,  Ahti’s  [1961])  treatment. 

The  keys  have  been  somewhat  expanded  to  include  brief  diagnoses 
of  each  species.  In  many  cases,  however,  additional  descriptive  com¬ 
ments  concerning  certain  important  or  confusing  taxa  have  been  included 
in  the  annotated  list. 

All  generic  and  specific  names  are  numbered  consecutively  in  the 
order  in  which  the  names  appear  in  the  annotated  list.  The  generic  keys 
following  the  group  keys  are  in  the  same  order.  References  to  the  generic 
keys  are  made  in  the  group  keys  using  the  genus  numbers  (in  boldface 
type).  All  genera  represented  by  only  one  species  on  Long  Island  do  not 
appear  in  the  generic  keys.  Instead,  the  species  is  keyed  out  in  the  group 
keys  and  is  directly  referred  to  its  position  in  the  annotated  list  by  means 
of  its  species  number  (in  lightface  type),  as  are  all  species  in  the  keys. 

Author  abbreviations  follow  Sayre,  et  al  (1964). 


112 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


In  the  annotated  list,  all  specimens  listed  under  “material  seen”  or 
elsewhere  in  the  discussions  which  were  collected  by  Imshaug  or  Brodo 
have  been  deposited  in  the  Michigan  State  University  Herbarium  (MSC), 
unless  otherwise  noted.  The  locations  of  other  specimens  have  been 
recorded  using  standard  abbreviations  (Lanjouw  and  Stafleu,  1964). 

Comments  on  extra-Long  Island,  North  American,  and  worldwide 
distributions  were  made  to  provide  a  framework  for  the  floristic  treat¬ 
ments  presented  in  the  section  on  Floristic  Elements.  No  attempt  was 
made  to  compile  a  complete  listing  of  all  known  localities.  Instead,  a 
limited  number  of  fairly  reliable  and,  in  most  cases,  modern  treatments 
were  consulted  to  provide  information  on  the  basic  distribution  patterns 
and  affinities.  Undoubtedly  some  of  the  records  are  based  on  old  concepts 
or  misidentifications  and  are  incorrect;  I  hope  that  the  errors  are  few. 

Individual  state  and  province  records  are  presented  where  specific 
statements  or  maps  of  distributions  are  not  available.  Sources  of  the 
North  American  records,  unless  stated  otherwise,  are  as  follows:  Nova 
Scotia  (Lamb,  1950);  Maine  ( Degelius,  1940;  Davis,  1964a,  b);  Con¬ 
necticut  (Evans  and  Meyrowitz,  1926;  Hale,  1950);  Massachusetts 
(Ahmadjian,  1958;  pers.  coll.);  New  Jersey  (pers.  coll.);  central 
New  York  (Brodo,  1959);  North  Carolina  (Degelius,  1941;  Culberson, 
1958a);  Tennessee  (Degelius,  1941;  Phillips,  1963):  Alabama  (McCul¬ 
lough,  1964);  Arkansas,  Missouri  (Hale,  1957b);  Oklahoma  (Hale,  1957b; 
Thomson,  1961);  Indiana  (Fink  and  Fuson,  1919);  Arizona  ( Darrow, 
1950;  Weber,  1963);  New  Mexico  (Rudolph,  1953b);  Michigan  (Hedrick 
and  Lowe,  1936;  Thomson,  1951);  Wisconsin  (Hale,  1955a;  Culberson, 
1955a);  Minnesota  (Fink,  1910);  Black  Hills  (Wetmore,  1965); 
Idaho  (Hedrick,  1948);  Washington  (Howard,  1950);  British  Columbia 
(Weber  and  Shushan,  1959);  Alaska  (Cummings,  1910;  Thomson,  1950b; 
Krog,  1962);  Northern  Saskatchewan  (Thomson  and  Scotter,  1961);  Mani¬ 
toba  (Thomson,  1953);  Ontario  (Thomson,  1955;  Ahti,  1964);  Quebec 
(Thomson,  1955);  Baffin  Island  (Hale,  1954);  Canadian  archipelago  and 
East  Arctic  (Thomson,  1960;  Lynge,  1935,  1947). 

LJnless  otherwise  stated,  European  records  are  based  on  Grummann 
(1963),  Poelt  (1963),  or  Zahlbruckner  (1922-40).  Statements  concern¬ 
ing  circumboreal  distributions  are  based  on  papers  by  Lynge  (1928, 
1938,  1940a,  1940b,  1940c),  as  well  as  the  papers  on  the  North  Ameri¬ 
can  arctic  cited  above.  Asian  references  are  all  presented  directly  in  the 
distributional  notes. 

Species  regarded  as  “endemic”  are  found  only  in  North  America, 
except  for  a  few  species  also  found  in  the  West  Indies. 


KEY  TO  GROUPS 

I.  Thallus  crustose:  attached  to  substrate  at  all  points;  lower  cortex 
absent  (if  podetioid,  see  Group  III;  if  squamulose,  see  Group  II) 
. 2 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  113 

1.  Thalius  at  least  partially  free  from  substrate . 3 

2.  Thalius  bearing  ascocarps . Group  I 

2.  Thalius  lacking  ascocarps . Group  II 


3.  Thalius  foliose:  lobes  flattened,  usually  broad,  clearly  dorsi-ventral, 
attached  to  substrate  either  directly  or  by  means  of  rhizines,  or 
rarely,  only  by  a  central  umbilicus;  lower  cortex  usually  present; 
apothecia  sessile  or  immersed,  thalius  never  podetioid.  .  .  .Group  III 
3.  Thalius  fruticose:  lobes  more  or  less  terete,  or  less  frequently,  flat¬ 
tened;  basally  attached  to  substrate  at  one  or  several  points;  pendu¬ 
lous,  caespitose,  or  podetioid . Group  IV 


GROUP  I  -  CRUSTOSE  LICHENS  (FERTILE  MATERIAL) 

1.  Phycobionts  blue-green  algae.  Thalius  dark  brown  to  black,  areolate 
to  subsquamulose,  isidiate,  prothailus  blue  green  or  blue  black;  apo¬ 
thecia  lecideine;  saxicolous  on  concrete.  .  .  .37.  Placynthium  nigrum 

1.  Phycobionts  green  algae . 2 

2.  Ascocarps  on  short,  hair-like  stalks;  hymenium  disintegrating  and 
spores  forming  a  yellow-  to  deep-brown  mazaedium.  Spores 

brown  (in  water),  spherical,  ca.  1-1.5  ij,  in  diameter . 

. 16.  Chaenotheca  phaeocephala 


2.  Ascocarp  sessile  or  immersed:  hymenium  remaining  intact.  .  .  .3 

3.  Ascocarp  ±  elongated;  irregular  or  obling . 4 

3.  Ascocarp  disk-shaped,  hemispherical  or  spherical  (sometimes  im¬ 
bedded  within  a  stroma) . 8 


4.  Spores  nonseptate,  hyaline,  3-4  x  7-13  pi,.  Lirellae  dark  brown  to 
red  brown  or  black,  oblong  or  elongate,  rarely  branched,  0.2  x 
0.35-0.55  mm . 28.  Xylographa  opegraphella 


4.  Spores  1  to  7  septate . 5 

5.  Spores  with  cylindrical  cells;  ascocarp  ascolocular . 6 

5.  Spores  with  lenticular  cells;  ascocarp  ascohymenial . 7 


6.  Ascocarp  ±  enclosed  in  a  heavy  carbonaceous  stromatic  wall .  . 

. 7.  Opegrapha 

6.  Ascocarp  without  a  carbonaceous  stroma  or  excipuloid  margin 

. 4.  Arthonia 

7.  Spores  hyaline,  5  to  7  septate,  32-48  x  6-9  ij.;  exciple  not  continuous 
below,  but  well  developed  laterally  and  projecting  conspicuously 

above  hymenium . 29.  Graphis  scripta 

7.  Spores  brown,  2  to  3  septate,  21-30  x  6-7  pi;  exciple  continuous  below, 

shallow,  i.e.,  not  projecting  appreciably  above  hymenium . 

. 30.  Phaeographis  dendritica 

8.  Ascocarp  spherical  or  flask  shaped,  with  walls  completely  enclos¬ 
ing  hymenium  except  for  ostiole  at  apex;  walls  generally  partly 

or  entirely  carbonaceous . 9 

8.  Ascocarp  disk  shaped  or  cup  shaped,  with  exposed  hymenium; 
or,  hymenium  enclosed  within  thalline  tissue  in  a  wart-like 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


1  14 


structure  as  in  Pertusaria,  without  carbonaceous  walls  ot 

any  kind . . ..16 

9.  Ascocarps  clustered  in  stromatic  verrucae,  more  than  one  per 

stroma . . 10 

9.  Ascocarps  single,  scattered . . 11 

10.  Thallus,  especially  stromatic  verrucae,  covered  with  a  rusty-red 
pigment  which  is  KOH  +  purple.  (Spores  brown,  3-septate,  but 
not  seen  in  L.I.  material).  Very  rare.  .  .  .23.  Melanotheca  cruenta 
10.  Thallus  brownish  or  olivaceous,  smooth,  KOH  ;  spores  hyaline, 
4  to  8  septate,  23-45  x  8-13  jx.  Frequent  on  Ilex  and  Fagus .  .  .  . 

. . . . . 24.  Trypethelium  virens 

11.  Spores  muriform,  hyaline.  Ascocarp  ascolocular . . . 

. . . .  .  4.  Polyblastiopsis  quercicola 


1  1.  Spores  non-septate  or  only  transversely  septate . .  12 

12.  Ascocarp  ascolocular . 13 

12.  Ascocarp  ascohymenial . . . . . 14 

13.  Spores  ellipsoid  or  fusiform,  straight,  15-21  x  (4-) 5-7  jx,  1  to  3 
septate;  on  oak  and  beach  plum . 1.  Arthopyrenia 


13.  Spores  acicular,  curved  or  sigmoid,  20-30  x  3-4  ix,  1  to  3  (-5)  septate; 

on  birch . .  3.  Leptorhaphis  epidermidis 

14.  Spores  brown,  3-septate,  cells  lenticular,  1  6-20 ( -25 )  x  1 0-1 3[x 

. . . . . 22.  Pyrenula  nitida 

14.  Spores  hyaline.  . . . . 15 

15.  Spores  nonseptate.  Saxicolous . .  .9.  Verrucaria 

15.  Spores  3  to  16  septate,  cells  cylindrical.  Corticolous.  ....  14.  Porina 

16.  Spores  more  than  50  per  ascus,  4  x  2  |x.  Saxicolous.  ........  17 

16.  Spores  ( 1  -)  8  ( -20 )  per  ascus,  usually  larger  than  4  x  2[x  .  ...  18 

17.  Thallus  epilithic,  areolate  to  squamulose,  brown;  apothecia  completely 

immersed  in  thallus.  . . .  141.  Acarospora  fuscata 

17.  Thallus  mostly  endolithic;  apothecia  sessile  with  a  lecideine  margin 
. . . .  .37.  Sarcogyne 


18.  Spores  muriform . .  .  .  . . 19 

18.  Spores  nonseptate  or  transversely  septate.  ...............  21 


19.  Corticolous.  Thallus  thin,  hypophloedal;  apothecia  minute,  puncti- 
form,  0.1 -0.2  mm  across;  spores  hyaline,  32-46  (-55)  x  10-23  (-27)  jx. 
Rare . . . . . 11.  Arthothelium  taediosum 


3  9.  Not  corticolous.  Thallus  well  developed;  apothecia  usually  larger  than 

0.2  mm;  spores  brown  or  sometimes  hyaline . .  20 

20.  Apothecia  deeply  concave,  imbedded  in  thick  thalline  verrucae 
resulting  in  a  double  margin  (thalline  and  proper);  spores  without 
any  gelatinous  epispore  (“halo”),  22-40  x  10-14  [x.  Medulla  C  + 
red,  KOH  +  yellow.  Saxicolous  or  growing  on  Cladonia.  Rare 

. . . .31.  Diploschistes  scruposus 

20.  Apothecia  flat  to  convex  with  proper  margin  alone,  imbedded  in 
thalline  verrucae  or  arising  between  them;  spores  with  a  gel  a- 


to  to  (O  to  to  to 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


115 


tinous  epispore  (“halo”).  Medulla  C  +  red  or  C KOH  +  or 

Saxicolous.  Common . 31.  Rhizocarpon 

1 .  Spores  brown,  uniseptate . 22 

1.  Spores  hyaline . 23 

22.  Apothecia  with  thalline  margin . 61.  Rinodina 

22.  Apothecia  without  thalline  margin . 60.  Biiellia 

3.  Apothecia  with  thalline  margin  or  enclosed  in  thalline  verrucae .  .24 

3.  Apothecia  without  thalline  margin . 31 

24.  Spores  vermiform  or  sigmoid,  septate  or  nonseptate,  length  to 
width  ratio  7-9:1,  45-62  x  5-8  [j..  Thallus  PD  +  orange  and 
KOH  +  yellow  (thamnolic  acid) . 


173.  Haematomma  ochrophaeum 
24.  Spores  ellipsoid,  oblong,  or  subspherical,  length  to  width  ratio 

ca.  1. 5-3:1 . 25 

.  Spores  over  40  tj,  long,  nonseptate . 26 

5.  Spores  under  30  [j.  long,  nonseptate  or  septate . 28 

26.  Apothecia  usually  imbedded  in  thalline  verrucae,  or,  if  lecanorine, 

then  spores  over  200  ;j.  long . 27 

26.  Apothecia  lecanorine;  spores  40-68  ij.  long,  8  per  ascus.  Disks 

C+  red . 43.  Ochrolechia 

27.  Spores  all  hyaline,  KOH  —  ;  spore  walls  not  radiately  channelled. 

Common . 39.  Pertusaria 


27.  Spores  sometimes  brownish,  KOH  +  sordid  violet,  125-190  x  30-45  jj.; 

spore  walls  conspicuously  channelled.  Uncommon . 

. 151.  Melanaria  macounii 

28.  Spores  polari'ocular.  Disk  KOH  +  red-violet  or  KOH— . 


. 57.  Caloplaca 

28.  Spores  nonseptate.  Disk  KOH- . 29 

29.  Apothecial  disk  and  margin  yellow.  Saxicolous.  .  .45.  CandelarieWa 


29.  Apothecial  disk  black,  brown,  pale  reddish  buff,  or  yellowish.  Saxico¬ 
lous  or  corticolous.  (If  disk  is  yellowish,  then  corticolous) . 30 

30.  Phycobiont  Trentepohlia;  apothecia  immersed  in  thallus.  Spores 

hyaline,  ellipsoid,  11-16  x  5-8  ij..  Saxicolous . 

. 152.  Ionaspis  odora 

30.  Phycobiont  Trebouxia;  apothecia  immersed  in  thallus  or  sessile 

. 42.  Lee  anora 

31.  Ascocarp  ascolocular.  Spores  usually  septate,  hyaline,  ellipsoid  to 


fusiform . 6.  Micarea 

31.  Ascocarp  ascohymenial . 32 

32.  Spores  septate . 33 

32.  Spores  nonseptate . 28.  Lecidea 

33.  Spores  uniseptate . 34 

33.  Spores  3  or  more  septate,  fusiform  to  acicular . 30.  Bacidia 

34.  Spores  polarilocular,  13-16  x  8-10  rj, .  Disks  KOH  +  purple-red 

. 227 .  Caloplaca  discolor 

34  Spores  not  polarilocular . 35 


1  16  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

35.  Apothecia  black,  strongly  convex  to  hemispherical;  hypothecium  dark 

brown;  spores  with  cells  of  unequal  size,  9-15  x  4-5  tj, . 

. 63.  Catillaria  glauconigrans 

35.  Apothecia  pale  pinkish  yellow  or  orange,  deeply  concave  to  ±  flat; 
hypothecium  hyaline;  spores  with  cells  of  equal  size,  9-14  x  2-4  p.. 

Asci  extremely  narrow,  almost  linear,  thin  walled . 36 

36.  Apothecial  disks  pink-yellow  (flesh  colored),  deeply  concave.  . 

. 32.  Dimerella  diluta 

36.  Apothecial  disks  pale  orange  to  orange  buff,  flat . 

. 33.  Dimerella  lutea 


GROUP  II  -  CRUSTOSE  LICHENS  (STERILE  MATERIAL) 

1.  Terricolous . 2 

1.  Saxicolous . 5 

1.  Corticolous  or  lignicolous . 15 

2.  Thallus  black  to  dark  brown,  minutely  verrucose  to  granulose. 

Thallus  KOH  -,  PD  -,  C  - . 59.  Lecidea  uliginosa 

2.  Thallus  pale  grey,  grey-green,  or  white . 3 

3.  Thallus  C  +  red,  KOH  -,  PD  - . 54.  Lecidea  granulosa 

3.  Thallus  C- . . 4 

4.  Thallus  PD  +  deep  yellow  (baeomycic  acid) . 

. 80.  Baeomyces  roseus 

4.  Thallus  PD -,  KOH  +  yellow  (atranorin) . 

. 78.  Pycnothelia  papillaria 

5.  On  calcareous  rock  or  mortar . 6 

5.  On  siliceous  rock . 7 

6.  Thallus  dark  brown  to  black,  isidiate,  KOH  -;  phycobionts  blue- 

green  algae.  Prothallus  conspicuous,  blue-green . 

. 37.  Placynthium  nigrum 

6.  Thallus  yellow  or  orange,  KOH  +  dark  purple;  phycobionts 
green  algae.  Thallus  granular  to  thickly  areolate  and  only  occa¬ 
sionally  breaking  into  sorediate  patches;  margin  of  thallus  diffuse 

. 226.  Caloplaca  citrina 

7.  Thallus  yellow  or  yellow-green,  KOH  - . 8 

7.  Thallus  white,  grey,  or  brown  (no  yellowish  tint)  .  . . 9 

8.  Thallus  margin  effigurate;  thallus  yellow-green.  Medulla  C  + 
red,  usnic  and  gyrophoric  acids  present.  .  .245.  Rinodina  oreina 

8.  Thalli  small,  scattered,  areolate  to  subsquamulose,  deep  yellow. 

Medulla  C  -,  usnic  and  gyrophoric  acids  absent . 

. 176.  Candelariella  vitellina 

9.  Medulla  C  +  red . 10 

9.  Medulla  C  - . 12 

10.  Thallus  grey,  smooth,  with  scattered  patches  of  soredia.  Medulla 
KOH- . (unknown  no.  I)7 


TThese  unidentified  sterile  crustose  species  have  been  deposited  in  herb.  MSC 
for  future  reference. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


117 


10.  Thallus  grey  to  ashy,  esorediate,  verrucose  to  areolate.  Medulla 
KOH  +  yellow  to  orange  and  PD  +  orange  (stictic  acid)  ....  11 

11.  Thallus  hrown,  verrucose.  Medulla  1  +  blue . 

. 74.  Rhizocarpon  grande 

11.  Thallus  grey  to  ashy,  areolate  to  verrucose.  Medulla  I- . 

. 75.  Rhizocarpon  intermedium 

12.  Thallus  leprose  to  granular  sorediate,  marginate  and  often  zoned. 
Thallus  PD  +  red  or  yellow,  KOH-  or  +  yellow)?)  (fumar- 

protocetraric  or  barbatolic  acid  present) . 

. 260.  Lepraria  zonata 

12.  Thallus  smooth  to  areolate  or  verrucose.  Thallus  medulla  KOH 

+  yellow  or  red  (stictic  or  norstictic  acid  present) . 13 

13.  Thallus  dark  cinereous  or  sordid  green-grey,  verrucose  to  areolate. 

Stictic  or  norstictic  acid  present . 157.  Lecanora  cinerea 

13.  Thallus  white  to  very  pale  grey  or  ashy,  smooth  to  areolate . 14 

14.  Prothallus  white,  often  conspicuous.  Stictic  or  norstictic  acid 

present.  Growing  on  stones  or  boulders  in  shaded  woods . 

. 48.  Lecidea  alhocaendescens 

14.  Prothallus  black,  often  conspicuous.  Norstictic  acid  present. 
Growing  on  exposed  boulders.  Pycnoconidia  short,  straight, 

bacilliform,  4-6  x  ca.  1  u. . 239.  Bnellia  stigmaea 

15.  Thallus  squamulose,  margins  entire.  Undersurface  of  squamules 

sorediate . 16 

(also  see  Cladonia  key) 

15.  Thallus  continuous  or  diffuse  (not  squamulose) . 17 

16.  Thallus  PD  +  red  ( fumarprotocetraric  acid),  C -.  Squamules 
dark  green-brown  to  olivaceous,  0.5-0.75  (-1.0)  mm  broad.  .  .  . 

. 49.  Lecidea  anthracophila 

16.  Thallus  PD -,  C  +  red.  Squamules  pale  olivaceous,  1.0-  1.5  mm 

broad . 58.  Lecidea  scalaris 

17.  Thallus  leprose,  sorediate,  or  coralloid-isidiate . 18 

17.  Thallus  smooth,  areolate,  or  verrucose . 32 

18.  Thallus  orange,  yellow,  or  yellowish  green . 19 

18.  Thallus  grey,  grey-green,  brown,  olivaceous,  or  black . 21 

19.  Thallus  dark  yellow  to  orange,  KOH  +  dark  purple.  Thallus  smooth, 

becoming  coarsely  sorediate  in  patches . 227.  Caloplaca  discolor 

19.  Thallus  yellow  to  yellowish  green,  KOH  - . 20 

20.  Lignicolous,  on  decorticate  Chamaecy paris  stumps  in  bogs.  Thal¬ 
lus  diffuse,  leprose,  pale  yellowish  or  whitish  green . 

. 16.  Chaenotheca  phaeocephala 

20.  Corticolous.  Thallus  leprose-granular,  deep  yellow . 

. 1  77.  Candelaria  concolor 

21.  Medulla  KC  +  violet.  Thallus  dark  cinereous  to  grey-green;  verrucae 

erupting  into  white  sorediate  mounds . 143.  Pertusaria  amara 

21.  Medulla  KC  -  or  KC  +  red . 22 

22.  Thallus  effuse,  leprose,  or  coralloid-isidiate . 23 


1  18  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

22.  Thallus  with  ±  distinct  soralia  at  least  at  thallus  margin,  verru- 

cose  or  ±  continuous . 27 

23.  Thallus  coralloid-isidiate;  phycobiont  Trentepohlia . 

. 27.  Porina  nucula 

23.  Thallus  effuse,  leprose,  phycobionts  Trebouxioid . 24 

24.  Thallus  bluish  green  or  blue-grey,  KOH  +  yellow  (atranorin), 

PD or  rarely,  PD  +  red  (fumarprotocetraric  acid) . 

. 259.  Lepraria  incana 

24.  Thallus  whitish  green  or  dark  green  to  blackish  green . 25 

25.  Thallus  KOH  +  yellow  and  PD  +  orange  (atranorin  and  stictic  acid). 
Thallus  pale  to  whitish  green,  with  thick  white  prothalline  mat.  .  .  . 

. 261.  Lepraria  sp. 

25.  Thallus  KOH -,  PD- . 26 

26.  Thallus  coarsely  granular,  pale  green  to  brownish  green.  Lignico- 

lous.  Very  rare . 62.  Lecidea  viridescens 

26.  Thallus  finely  granular,  dark  green  to  hlackish  green.  Mostly 

corticolous.  Very  common . 66.  Bacidia  chlorococca 

27.  Thallus  composed  of  scattered  verrucae  or  areoles,  some  bursting 

into  soredia . 28 

27.  Thallus  ±  continuous  and  smooth,  at  least  at  the  margins . 29 

28.  Medulla  C  +  red . 47.  Lecidea  aeruginosa 

28.  Medulla  C- . 50.  Lecidea  botryosa 

29.  Thallus  KOH  +  deep  yellow  and  PD  +  orange  (thamnolic  acid)  .  . 

. 30 

29.  Thallus  KOH -,  PD- . 31 

30.  Thallus  pale  grey  to  white,  with  crowded  hollow  verrucae  in  the 
older  portions,  many  of  which  burst  revealing  coarsely  granular 
soredia  often  leaving  the  center  of  the  thallus  essentially  leprose 

. . . 174.  Haematomma  sp. 

30.  Thallus  ashy  or  darker,  with  sorediate  verrucae  scattered  evenly 

over  the  thallus . 147.  Pertusaria  trachythallina 

31.  Thallus  greenish  or  brownish  green  with  maculiform  greenish  or 

yellow-green  soralia  scattered  over  the  thin  thallus . 

. Cfr.  Opegrapha  sp. 

31.  Thallus  grey  or  greenish-grey,  bursting  into  scattered,  granular-soredi- 

ate  soralia.  Hypophloedal;  phycobiont  Trentepohlia . 

. (unknown  no.  6) 8 


32.  Medulla  KOH  +  yellow  or  red . 33 

32.  Medulla  KOH- . 35 


33.  Medulla  KOH  +  yellow.  Thallus  thick  or  thin,  pale  grey  to  dark  ashy 

. 34 

33.  Medulla  KOH  +  red  (norstictic  acid).  Thallus  thin,  smooth,  becom¬ 
ing  areolate  or  chinky,  pale  greenish  grey  to  white;  pycnoconidia  4-7  p, 

long,  straight,  bacilliform . 

. 235.  Buellia  curtisii  or  240.  B.  stillingiana 


"/hid 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  119 

34.  Medulla  KOH  +  deep  yellow  and  PD  +  orange  (thaninolic 

acid).  Thallus  densely  verrucose  and  rugose,  pale  grey . 

. 173.  Haematomma  ochrophaeum 

34.  Medulla  KOH  +  yellow-orange,  PD-  (?),  thaninolic  acid  and 
atranorin  absent,  (stictic  acid  present?)  thallus  very  thin,  smooth, 

greenish  grey.  Pycnoconidia  short,  straight,  4-5  x  ca.  1  tj . 

. (unknown  no.  7)9 

35.  Thallus  olivaceous  to  blackish  green,  well  developed,  rugose  to  verru- 

culose;  pycnidia  common,  brown,  pycnoconidia  0.5  x  1 .2  pc . 

. 65.  Bacidia  chlorantha 

35.  Thallus  very  thin  or  hypophloedal,  or,  if  thicker,  ashy  or  pale  green¬ 
ish  grey;  pycnidia  common,  black;  pycnoconidia  over  4  p.  long.  .  .  .36 

36.  Pycnidia  ±  clustered  in  small  groups;  pycnoconidia  4-5  x  1  p., 

straight,  bacilliform.  On  Ilex  and  Fagus . 

. 24.  Trypethelium  virens 

36.  Pycnidia  scattered  evenly  over  the  thallus;  pycnoconidia  curved 

. 37 

37.  Pycnoconidia  reniform,  short,  broad,  5-7  x  3-4  ij. . 

. 15.  Opegrapha  rufescens 

37.  Pycnoconidia  sickle  shaped,  slender  (1  tj,  broad) . 38 

38.  Pycnoconidia  10-15  p.  long  (measured  end  to  end,  in  a  straight 

line),  very  strongly  curved . 14.  Opegrapha  cinerea 

38.  Pycnoconidia  15-2()p.  long  (measured  as  above),  slightly  curved 
. ( unknown  no.  5) 9 

GROUP  III  -  FOLIOSE  LICHENS 

1.  Thallus  composed  of  aggregations  of  squamules  individually  attached 
to  the  substrate  at  one  edge;  (0.5-)  1-3  (-5)  mm  long  or  broad.  .  .  . 

. 35.  Cladon  ia 

1.  Thallus  centrally  attached;  squamules,  if  present,  part  of  a  broad 
thallus;  thallus  over  10  mm  in  diameter . 2 


2.  Phycobionts  blue-green  algae . 3 

2.  Phycobionts  green  algae . 7 

3.  Thallus  gelatinous  when  moistened . 4 

3.  Thallus  not  gelatinous  when  moistened . 5 


4.  Upper  cortex  absent;  globular  isidia  present;  thallus  broad.  .  .  . 

. 34.  Collema  sabfurvum 

4.  Upper  cortex  present,  paraplectenchymatous;  isidia  absent  or 

coralloid-cylindrical;  thallus  narrow  lobed . 21 .  Leptogium 

5.  Thallus  small,  lobes  2-3  mm  broad;  apothecia  scattered  over  the  sur¬ 
face  of  thallus;  spores  nonseptate.  Lower  surface  densely  white  or 

tan,  tomentose . 38 .  Pannaria  lurida 

5.  Thallus  large,  lobes  3-30  mm  broad;  apothecia  at  tips  of  lobes; 

spores  septate . 6 

6.  Lower  surface  ecorticate,  usually  conspicuously  veined;  apothecia 
on  upper  surface  of  lobes;  medulla  KOH-.  .  .27.  Peltigera  (p.p.) 


Ibid 


120 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


6.  Lower  surface  corticate,  glabrous,  without  veins;  apothecia  on 
lower  surface  of  lobes;  medulla  yellow,  KOH  +  pink  to  red- 
violet  (anthraquinone:  nephromin)  .  .  .41.  Nephroma  laevigatum 


7.  Thallus  bright  yellow  or  orange . 8 

7.  Thallus  brownish,  grey,  grey-green,  or  yellowish  green . 9 


8.  Upper  cortex  KOH  +  red-violet  (anthraquinone:  parietin)  .... 

. 58.  Xanthoria 

8.  Upper  cortex  KOH-  (pulvic  acid  derivative) . 

. 177.  Candelaria  concolor 

9.  Thallus  attached  to  the  substrate  by  central  umbilicus . 10 

9.  Thallus  attached  to  substrate  directly,  or  by  many  fine  rhizines  ....  12 

10.  Thallus  yellow-green.  Ascocarps  apothecia  with  orange  disks, 


abundant . 165.  Lecanora  rubina 

10.  Thallus  brown,  with  no  yellow  tint . 11 


11.  Ascocarps  (usually  present)  perithecia;  medulla  C- . 

. 21.  Dermatocarpon  miniatum 

11.  Ascocarps  (if  present)  apothecia  with  black  disks,  medulla  C  + 

red . 36.  Umbilicaria 

12.  Cephalodia  abundant,  scattered  over  the  upper  surface  of  thallus. 

Phycobiont  Coccomyxa.  Very  rare;  on  soil . 

. 43.  Peltigera  aphthosa 

12.  Cephalodia  absent . 13 

13.  Thallus  inflated,  hollow;  lower  surface  corticate,  brown  to  black, 
smooth,  naked.  Granular  soredia  in  labriform  soralia;  medulla  PD  + 
red  ( monoacetyl-protocetraric  acid),  KC  +  red  (physodic  acid)  .  .  . 

. 203.  Hypogymnia  physodes 

13.  Thallus  solid;  lower  surface  rhizinate,  tomentose,  or  ecorticate.  ...  14 

14.  Hypothallus  present,  composed  of  a  thick  mat  of  interwoven 
black  hyphae.  Medulla  PD-,  KOH-,  KC-.  .  .210 .  Anzia  colpodes 

14.  Hypothallus  lacking . 15 

15.  Lower  surface  feltlike  or  tomentose,  without  rhizines.  Lobes  broad, 

over  3  mm  across . 16 

15.  Lower  surface  rhizinate . 17 

16.  Medulla  PD-,  C-.  Apothecia  common,  immersed  in  depressions 
in  lobes;  spores  brown,  uniseptate,  4  per  ascus;  phycobiont 

Coccomyxa . 42.  Solorina  saccata 

16.  Medulla  PD  +  orange  (stictic  acid),  or,  C  +  red  (gyrophoric 
acid).  Apothecia,  if  present,  sessile;  spores  hyaline,  3-septate, 


8  per  ascus;  phycobiont  Trebouxia . 24.  Lobaria 

17.  Thallus  yellow  or  yellow-green . 18 

1 7.  Thallus  brown,  grey,  or  grey-green . 20 


18.  Lower  surface  bright  yellow;  usnic  acid  absent.  Thallus  smooth 
or  rugose;  soredia  and  isidia  absent;  black  pycnidia  common 
along  thallus  margins,  sometimes  becoming  partially  laminal. 

Medulla  PD-,  KOH-,  C-,  KC- . 209.  Cetraria  viridis 

18.  Lower  surface  not  yellow;  usnic  acid  present . 19 


to  K)  to  to  to  to 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  121 

19.  Thallus  with  lobes  less  than  1  mm  broad;  older  portions  covered  with 

granular  soredia;  divaricatic  acid  present . 

. 179.  Parmeliopsis  ambigua 

19.  Thallus  with  lobes  broader  than  1  mm;  soredia  present  or  absent; 

divaricatic  acid  absent . 48.  Parmelia  ( p.p. ) 

20.  Medulla  PD  +  orange.  KOH  +  deep  yellow  (thamnolic  acid)  .  . 

. 47.  Parmeliopsis  (p.p.) 

20.  Medulla  not  having  that  combination  of  reactions  (thamnolic 


acid  absent) . 21 

1.  Lower  surface  white,  pale  buff,  or  yellow . 22 

1.  Lower  surface  light  or  dark  brown,  or  black  (although  marginal  areas 

may  have  broad,  irregular,  white  blotches) . 26 

22.  Thallus  brown  or  olivaceous-brown.  Medulla  C  +  red . 

. 199.  Parmelia  subaurifera 

22.  Thallus  grey  or  grey-green . 23 

3.  Thallus  lobes  3-7  mm  broad . 24 

3.  Thallus  lobes  0.5-3  mm  broad . 25 

24.  Pseudocyphellae  on  upper  surface;  medulla  I- . 


. 48.  Parmelia  (p.p.) 

24.  Pseudocyphellae  absent;  medulla  I  +  blue . 

. 208.  Cetraria  tuckermanii 

.  Cortical  hyphae  parallel  with  surface . 64 .  Anaptychia 

5.  Cortical  hyphae  perpendicular  to  surface . 63 .  Physcia 

26.  Rhizines  black  with  white  tips,  very  dense;  lobes  1-2  mm  broad 

. 27 

26.  Rhizines  uniform  in  color,  sparse  to  dense;  lobes  1-6  mm  broad 

. 28 

27.  Medulla  mustard  yellow,  KOH  +  dull  red-brown;  lobes  pruinose, 

especially  near  tips,  with  granular  marginal  soredia . 

. 248.  Pyxine  sorediata 

27.  Medulla  red-orange  (KOH  +  purple)  or  white  (KOH-);  lobes  not 

pruinose;  soredia  marginal  and  laminal . 252.  Physcia  orbicularis 

28.  Pycnidia,  common,  marginal;  rhizines  sparse . 

. 51 .  Cetraria  (p.p.) 

28.  Pycnidia  rare,  laminal;  rhizines  usually  ±  dense . 

. 48.  Parmelia  (p.p.) 

GROUP  IV  —  FRUTICOSE  LICHENS 

1 .  Thallus  having  erect,  terete  or  subterete  podetia  or  pseudopodetia. 

Mostly  terricolous,  but  sometimes  corticolous  or  saxicolous . 2 

1.  Thallus  not  podetioid;  erect  and  shrubby,  or,  more  or  less  pendent. 


Corticolous  or  saxicolous . 5 

2.  Podetia  or  pseudopodetia  hollow . 3 

2.  Podetioid  structures  solid . 4 


3.  Primary  thallus  squamulose  or  soon  absent;  spores  nonseptate . 

. 35.  Cladonia 


122  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

3.  Primary  thallus  crustose,  persistent,  white  granular;  spores  uniseptate. 

Medulla  KOH  +  yellow  (atranorin) . 78.  Pycnothelia  papillaria 

4.  Primary  thallus  consisting  of  white  granules;  podetia  short,  each 
one  terminated  by  a  large,  pink  apothecium.  Podetia  and  thallus 
PD  +  yellow,  KC-  (baeomycic  acid).  On  raw,  eroding  soil.  .  .  . 

. . .  80.  Baeomyces  roseus 

4.  Primary  thallus  consisting  of  prong-like  phyllocladia;  pseudo- 
podetia  sterile  or  with  brown  apothecia.  Medulla  PD-,  KC  +  red 

(lobaric  acid).  Saxicolous . 79.  Stereocaulon  saxatile 

5.  Thallus  composed  of  terete  filaments . 6 

5.  Thallus  composed  of  distinctly  flattened  or  at  least  basally  angular 

lobes  or  branches . 7 

6.  Thallus  dark  brown;  filaments  having  no  central  cartilaginous 
axis.  Soralia  present.  Medulla  PD  +  red  ( fumarprotocetraric 

acid . .  54.  Alectoria 

6.  Thallus  yellow-green  to  grey-green;  filaments  with  a  central,  car¬ 
tilaginous,  elastic  axis . 56.  Usnea 

7.  Thallus  light  or  dark  brown,  shrubby;  terricolous.  Marginal  pycnidia 
abundant;  pseudocyphellae  linear,  submarginal; 

. 207.  Cetraria  islamlica  subsp.  crispa 

7.  Thallus  not  brown;  corticolous . 8 

8.  Thallus  yellow  or  orange.  Cortex  KOH  +  red-violet . 

. 59.  Teloschistes 

8.  Thallus  grey-green  or  yellow-green.  Cortex  K-  or  K  +  yellow  ...  9 

9.  Medulla  C  +  red.  Thallus  isidiate,  grey-green  (usnic  acid  absent); 

clearly  dorsi-ventral . 204.  Pseudevernia  furfuracea 

9.  Medulla  C-.  Thallus  not  isidiate,  yellow-green  to  grey-green  (usnic 

acid  present);  upper  and  lower  surfaces  not  distinguishable . 10 

10.  Thallus  soft,  flexible  (without  chondroid  layer),  sorediate.  Me¬ 
dulla  KOH-,  PD- . 211.  Evernia  mesomorpha 

10.  Thallus  stiff,  (with  chondroid  layer),  often  caespitose,  esorediate. 
Medulla  KOH-  or  KOH  +  red . 55.  Ramalina 

1.  ARTHOPYRENIA 

1.  Spores  ellipsoid  to  subfusiform,  15-17  x  5-7  p,,  1  to  3  septate,  with 
cells  usually  unequal  in  size;  pseudothecia  0.1 5-0.26  mm  in  diameter; 
paraphysoid  threads  persistent,  distinct.  Corticolous.  .  .2 .  A.  pinicola 
1.  Spores  fusiform,  16-21  x  4-5 (-7) 1  to  3  septate,  with  cells  equal 
in  size;  pseudothecia  0.15-0.25  mm  in  diameter;  paraphysoid  threads 
distinct  and  persistent.  Corticolous . 1  .A.cerasi 

4.  ARTHONIA 

1.  Phycobiont  Trebouxia.  Thallus  whitish  to  yellowish  green,  granular  to 
verrucose;  ascocarps  round;  disks  ashy  grey  to  black,  heavily  pruinose; 

spores  3-septate,  ( 1 4- )  1 6-22  x  5-7  g. . 5.  A.  caesia 

1.  Phycobiont  Trentepohlia . 2 

2.  Ascocarps  jet  black  or  bluish  grey  (even  when  moist) . 3 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


123 


2.  Ascocarps  red-brown  to  dark  brown  or  brownish  black,  turning 

a  distinct  red-brown  when  moistened . 5 

3.  Hypothecium  (fruit  base)  brown.  Thallus  scattered,  granulose  to  dis¬ 
appearing;  ascocarps  punctiform;  spores  3-septate  ±  clavate,  10-17  x 

4-6  p . 6.  A.  cfr.  mediella 

3.  Hypothecium  (fruit  base)  hyaline  or  essentially  absent . 4 

4.  Spores  (3-)  5  septate,  penultimate  cells  much  shorter  than  other 

cells,  1 7-20  x  5-7  p . 9.  A.  sexlocularis 

4.  Spores  3-septate.  all  spore  cells  equal  in  size,  14-20  x  (4-)5-7p 

. 8.  A.  punctiformis 

5.  Spores  2  to  4  septate,  hyaline,  one  end  cell  much  larger  than  other 

cells;  ascocarps  epruinose;  spores  14-20  x  5-7  p . 10.  A.  siderea 

5.  Spores  constantly  3-septate,  ashy  brown,  all  cells  equal  in  size;  asco¬ 
carps  heavily  pruinose;  spores  12-17  x  4-6  p . 1.  A.  polymorpha 

6.  MICAREA 

1 .  Saxico'.ous.  Thallus  greenish  or  brownish  grey,  minutely  verrucose  to 
granulose;  ascocarps  less  than  0.5  mm  in  diameter,  buff  to  black; 
spores  3-septate,  (8-)  12-16  x  (2-)3-4p.  .  .  .71.  Bacidia  cfr.  trisepta 

1.  Lignico’ous  (on  rotting  wood) . 2 

2.  Spores  mostly  uniseptate,  sometimes  nonseptate,  6. 5-8. 5  x  3.0- 
3.5  p;  thallus  blackish  green,  minutely  granulose;  ascocarps  very 

convex  to  hemispherical,  brown  to  black . 13.  M.  prasina 

2.  Spores  1  to  3  septate,  16-19  x  5-6  p;  thallus  dark  green  to  green¬ 
ish  black,  smooth  or  verrucose  to  ±  granulose;  ascocarps  very 
convex  to  hemispherical,  pitch  black . 12.  M.  melaeua 

7.  OPEGRAPHA 

I.  Thallus  thin,  continuous  to  scurfy  or  hypophloedal;  spores  8  per 

ascus . 2 

1.  Thallus  thin,  becoming  sorediate  in  maculiform  yellow  green  soralia; 
spores  4  per  ascus.  Ascocarps  short  and  broad,  0.5-0.65  x  0.11- 
0.4  mm,  somewhat  branches;  spores  5  to  7  septate,  18-23  x  4-5  p.  .  . 

. Cfr.  Opegrapha  sp. 

2.  Spores  22-36  x  2-3  p,  3  to  7  septate;  ascocarps  ( 0.25-) 0.5-2  mm 
long,  somewhat  branched,  pycnoconidia  9-15  x  1  -2  p,  strongly 

curved  or  twisted . 14.  O.  cinerea 

2.  Spores  19-24  x  3.5  p,  I  to  3  septate;  ascocarps  up  to  0.5  mm 

long,  unbranched;  pycnoconidia  5-7  x  1-2  p,  curved . 

. 15.  O.  rufescens 

9.  VERRUCARIA 

1.  Spores  6-9  x  3-5  p;  perithecia  0.1 -0.2  mm  across;  thallus  very  thin, 

filmy,  sordid  dark  brown.  On  quartz  pebbles  in  littoral  zone . 

. 17.  V.  microspora 

1.  Spores  15-26  x  6-15  p;  perithecia  0. 2-0.4  mm  across . 2 

2.  On  littoral  quartz  pebbles.  Thallus  smooth,  extremely  thin,  con¬ 
tinuous,  hlack  to  dark  brown;  spores  16-25  x  6-10  p . 


124 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


. 20.  V.  silicicola 

2.  On  concrete  and  mortar . 3 

3.  Thallus  thick,  dark  brown  to  brownish  grey,  dispersed  verrucose, 
areolate  to  almost  squamulose;  exciple  carbonaceous;  spores  15-18  x 

8-9  p, . 1  9.  V.  nigrescens 

3.  Thallus  thin,  pale  grey  to  whitish  ash,  areolate  to  chinky,  the  areoles 

being  =b  dispersed;  exciple  pale;  spores  20-23 ( -26 )  x  10-14p. . 

. 18.  F.  murdlis 


14.  PORINA 

1.  Perithecia,  buff  to  tan,  0.2-0. 3  mm  across;  spores  5  to  9  septate, 

48-75  x  7-9  [i.  Thallus  effuse  coralloid-isidiate;  exciple  pale . 

. 27.  P.  nucula 

1.  Perithecia  black;  spores  less  than  6.5  rj.  broad . 2 

2.  Spores  3  to  7  septate,  30-42  x5-6[x;  exciple  pale;  thallus  greenish 

black,  chinky  to  almost  granulose,  well  developed . 

. 25.  P.  cestrensis 

2.  Spores  mostly  9  to  13  septate,  58-65  x  5-7  p.;  exciple  carbonace¬ 
ous;  thallus  dark  or  light  grey-green,  diffuse,  very  thin,  almost 


absent  in  places . 26.  P.  hibernica 

21.  LEPTOGIUM 

1.  Thallus  very  thin,  isidiate,  the  isidia  cylindrical,  becoming  coralloid 

and  subsquamulose;  apothecia  absent . 36 .  L.  cyanescens 

1.  Thallus  relatively  thick,  not  isidiate  or  sorediate,  but  rugose  and  finely 
rugulose;  apothecia  common;  margins  smooth  and  entire;  spores 
20-23  x  9-12  tj, . 35.  L.  corticola 

24.  LOBARIA 


1.  Thallus  olivaceous,  pitted  and  reticulate,  with  soredia  and  sometimes 
isidia  on  the  ridges  and  margins.  Sterile  on  L.I.  Medulla  PD  +  orange 
and  KOH  +  yellow  (stictic  acid),  C-,  KC  +  reddish  (lobaric  acid?) 

. 39.  L.  puhnonaria 

1 .  Thallus  grey  to  light  green,  smooth,  without  soredia  or  isidia.  Usually 
fertile.  Medulla  PD-,  KOH-,  C  +  red . 40.  L.  quercizans 


27.  PELTIGERA 

1 .  Phycobionts  green  algae,  cephalodia  scattered  over  thallus  surface. 


Rare . 43.  P.  aphthosa 

1.  Phycobionts  blue-green  algae;  cephalodia  absent . 2 

2.  Thallus  surface  glabrous  (without  tomentum).  Spores  acicular 

75-103  x  4-5  p, . 45.  P.  polydactyla 

2.  Thallus  surface  tomentose  to  some  extent . 3 

3.  Thallus  producing  minute  regeneration  squamules  at  edges  and  along 

wounds . 46.  P.  praetextata 

3.  Thallus  not  producing  regeneration  squamules.  .  .  .44.  ( P.canina )  4 

4.  Thallus  with  grey  granular  soredia  produced  in  small,  laminal, 

orbicular  soralia . 44.  P.  canina  var.  spuria 

4.  Thallus  esorediate . 5 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


125 


5.  Veins  on  lower  surface  white . 

5.  Veins  brown  to  the  edge  of  the  thallus . 


.  44.  P.  canina  var.  rufescens 
.  44.  P.  canina  var.  ulorrhiza 


28.  LECiDEA 

1 .  On  soil . . . 2 

1.  On  rock . . . 3 

1.  On  bark  or  old  wood . 8 

2.  Thallus  green  to  greenish  grey  or  greenish  white,  verrucose, 
becoming  sorediate,  C  +  red;  apothecia  0.6-1. 3  mm  in  diameter; 

hypothecium  hyaline;  spores  6-10  x  3-6  ij, . 54.  L.  granulosa 

2.  Thallus  dark  olivaceous  brown  to  black,  granulose,  C-;  apothecia 
mostly  0. 3-0.4  mm  in  diameter;  hypothecium  dark  brown;  spores 

( 6-)  8-10  x  4-7  [a . 59.  L.  uliginosa 

3.  Apothecia  white  pruinose  with  conspicuous  dark  grey  rims.  Thallus 
light  grey  to  whitish  grey,  continuous  to  irregularly  cracked,  KOH  + 
red  (norstictic  acid)  or  KOH  +  yellow  (stictic  acid);  hypothecium 
dark  brown  or  red-brown;  spores  (13-)  16-20  x  6-8 (-10)  [a.  Usually  on 

shaded  rocks . 48.  L.  albocaerulescens 

3.  Apothecia  black  or  brown,  epruinose.  Thallus  KOH- . 4 

4.  Apothecia  0.5-1. 5  mm  in  diameter;  disks  black . 5 

4.  Apothecia  less  than  0.5  mm  in  diameter;  disks  black  or  brown ...  6 

5.  Spores  16-18  x  8  [j, . 55 .  L.  macrocarpa 

5.  Spores  7-12  x  3-6  p, . 52.  L.  cfr.  cyrtidia 

6.  Spores  11-20  x  7-10  ij.;  apothecial  disks  red-brown  to  dark  brown 

to  black;  hypothecium  yellowish  to  hyaline . 51.  L.  coarctata 

6.  Spores  6-8  x  3-4  p,;  apothecial  disks  black;  hypothecium  dark 

brown . 7 

7.  Epithecium  and  outer  edge  of  exciple  dark  green  to  greenish  black .  . 

. 53.  L.  erratica 

7.  Epithecium  and  outer  edge  of  exciple  reddish  brown,  not  green  (but 

hymenium  may  be  pale  olivaceous  at  times) . 52.  L.  cyrtidia 

8.  Thallus  squanuilose,  composed  of  imbricate  squamules.  Apo¬ 
thecia  rare . 9 

8.  Thallus  not  squamulose . 10 

9.  Thallus  C  +  red,  PD-  (lecanoric  acid).  Squamules  mostly  0. 5-1.0 

mm  across,  yellowish-  or  olive-green . 58.  L.  scalaris 

9.  Thallus  C-,  PD  +  red  (fumarprotocetraric  acid).  Squamules  mostly 
less  than  0.5  mm  across,  olive-  to  brownish  green  to  dark  olivaceous 

brown . . . 49.  L.  anthracophila 

10.  Hypothecium  dark  brown  or  reddish  brown . 11 

10.  Hypothecium  hyaline . 12 

11.  Thallus  thick  verrucose-areolate,  becoming  sorediate,  grey-green  to 

brown;  spores  6-12  x  (2-)3-5[j, . 50 .  L.  botryosa 

1  1.  Thallus  not  sorediate,  very  thin,  dark  green-black;  spores  6-8  x  3-4  [j. 

. 56.  L.  myriocarpoides 

12.  Spores  narrowly  ellipsoid  to  fusiform,  11-19  x  3-5  pi . 13 


126 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


12.  Spores  ellipsoid  to  spherical,  5=10  x  3-7  p.  Apothecial  disks  red- 

brown  to  black . . . . . . 14 

13.  Spores  15-19  x  4-5  p,  sometimes  uniseptate;  apothecia  often  strongly 
convex  and  hemispherical,  disks  flesh  colored  to  darker  brown .... 

. . . .  61.  L.  vernalis 

13.  Spores  11-13  x  3-4  p,  never  uniseptate;  apothecia  ± convex  but  not 

hemispherical,  disks  yellow  to  pale  orange . 

. (see  167.  Lecanora  symmicta) 

14.  Thalius  yellow-green,  areolate  to  chinky  or  somewhat  granular, 
C  +  yellow-orange.  Apothecia  red-brown  to  dark  brown,  usually 
less  than  0.3  mm  across;  spores  7-10x  (4-)5-7  p .  .  .  60.  L.  various 

14.  Thalius  grey-green  to  brownish  green,  granulose  to  sorediate, 

C  +  red  or  C- . . . . . 15 

15.  Spores  subglobose,  5-6  x  3-6  p.  Soredia  C- .  .  .  . . 57.  L.  nylanderi 

15.  Spores  ellipsoid  or  oval,  6-10  x  3-4 jx.  Soredia  C  +  red . 16 

16.  Apothecial  disks  lead  black,  margins  prominent . 

. . . 47.  L,  aeruginosa 

16.  Apothecial  disks  black,  margins  absent.  ....  .62.  L.  viridescens 

30.  BACIDIA 

1.  Spores  narrowly  ellipsoid  or  narrowly  ovate;  ratio  of  length  to  width 


not  more  than  7:1 . 2 

1.  Spores  acicular,  very  narrow,  ratio  of  length  to  width  usually  more 

than  7:1 . 4 

2.  Saxicolous.  Apothecia  minute,  pale  buff  to  black,  convex,  margin¬ 
less;  spores  (8-)  12-16  x  ( 2-)  3-4  [x;  thalius  coarsely  granulose, 

light  brownish  grey  to  greenish  grey . 71.fi.  cfr.  trisepta 

2.  Corticolous . 3 


3.  Hypothecium  dark  brown;  hymen ium  brownish;  spores  16-20  x  5-6  p 

. . . . . (see  12.  Micarea  melaena ) 

3.  Hypothecium  and  hymenium  hyaline;  spores  (1 9-)  23-32  x  3-6  [x. 
Apothecia  minute,  black,  convex,  marginless.  .66 .  Bacidia  chlorococca 

4.  Spores  strongly  curved  and  spiral  shaped,  13-16  x  2-3  [x  (mea¬ 
sured  end  to  end,  in  a  straight  line).  Saxicolous.  Rare. . 

. . . 72.  B,  umbrina 

4.  Spores  ±  straight . . . 5 

5.  Saxicolous.  Disks  usually  lighter  than  margins;  epithecium  dark 
greenish  black  to  black;  spores  obscurely  3-septate,  19-28  x  1-2  p 

. . . .  . . .  69.  B.  inundata 

5.  Corticolous  . . . . . . 6 

6.  Hypothecium  dark  brown  or  red-brown . . . 7 

6.  Hypothecium  pale,  hyaline,  yellowish,  or  very  light  brown ....  8 

7.  Apothecia  small,  0.25-0.60  mm  in  diameter;  spores  obscurely  1  to  3 
septate,  20-33  x  2-3  p;  phycobiont  Trebouxia .  .  .  .  .67.  B.  chlorosticta 

7.  Apothecia  large,  0.75-1.25  mm  in  diameter;  spores  obscurely  6  to  9 
septate,  35-55  x  3-4  p;  phycobiont  Trentepohlia .  .  .70.  B.  schweinitzii 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


127 


8.  Polysporous;  thallus  thick,  coarsely  granular  to  verrucose,  dark 
green  to  olive;  apothecia  irregular,  up  to  1.25  mm  in  diameter, 
often  clustered  and  crowded;  margins  raised,  conspicuous.  Spores 

5  to  7  septate,  20-26  x  2-3  p . 65.  B.  chlorantha 

8.  Octosporous;  thallus  thin,  greenish  grey;  apothecia  smaller, 

round,  not  clustered;  margins  not  raised . 9 

9.  Disks  red-brown  to  black;  margins  concolorous  or  lighter,  disappear¬ 
ing  with  age;  epithecium  reddish  violet  (intense  in  KOH);  spores  7  to 

many  septate,  39-68  x  4-6  jj. . 64 .  B.  atrogrisea 

9.  Disks  light  buff  to  ±  dark  brown  (never  black);  margins  darker 
than  disks;  epithecium  brown;  spores  obscurely  3  to  4  septate,  19-32 
x  l-3u, . 68.  B.  intermedia 

31.  RHIZOCARPON 

1.  Spores  uniseptate,  hyaline  to  slightly  tinted,  ( 1 1  -)  1 3-20  x  (5-)6-10[jl 
Thallus  sordid  greyish  green  to  ashy,  verrucose  to  minutely  verruculose, 
KOH  +  red  (norstictic  acid)  or  KOH  +  yellow  (stictic  acid)  .... 

. 73.  R.  cinereovirens 

1.  Spores  muriform  or  submuriform,  or  thallus  sterile . 2 

2.  Medulla  C  +  red  (gyrophoric  acid?),  KOH  +  yellow  (stictic 

acid).  Spores  soon  dark  brown,  many  celled . 3 

2.  Medulla  C-,  KOH  +  red  or  KOH-.  Spores  hyaline  for  a  long 

time,  then  brown . 4 

3.  Medulla  I-.  Thallus  whitish  to  light  ashy  or  brownish  grey,  subcon- 
tinuous  to  areolate,  and  finally  verrucose;  spores  26-38  x  10-15  p, .  . 

. 75.  R.  intermedium 

3.  Medulla  I  +  blue.  Thallus  dark  brown  or  grey  brown,  verrucose  with 

±  round,  ±  scattered  verrucae;  spores  25-29  x  10-13  p . 

. 74.  R.  grande 

4.  Medulla  KOH  +  red  (norstictic  acid).  Thallus  ±  smooth,  thin; 
apothecia  without  any  indication  of  a  thalline  margin;  spores 

20-27  x  10-13  n . 77.  R.  plicatile 

4.  Medulla  KOH-.  Thallus  verrucose  or  areolate,  almost  squamulose 
in  places;  apothecia  immersed  in  small  areoles  giving  appearance 

of  a  thalline  margin;  spores  1 9-29 (-32)  x  8-16  p . 

. 76.  R.  obscuratum 

35.  CLADONIA 

1 .  Primary  thallus  crustose,  persistent,  consisting  of  grey-green  or  grey 
to  whitish  verrucae  or  granules;  spores  uniseptate.  Pseudopodetia 
usually  under  0.75  mm  tall,  molariform  to  somewhat  branched,  often 

inflated.  Pseudopodetia  KOH  +  yellow  and  PD-  (atranorin) . 

. 78.  Pycnothelia  papillaria 

1.  Primary  thallus  squamulose  or  absent  in  mature  plant;  spores  non- 

septate . 2 

2.  Podetia  forming  a  more  or  less  complex  branch  system  (shrubby) ; 
primary  thallus  disappearing  in  maturity . 3 


128 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


2.  Podetia  simple  or  sparingly  branched,  or  absent ,  primary  thallus 

squamulose,  persistent . 16 

3.  Podetia  corticate,  except  where  cortex  is  replaced  by  soredia  in  the 

sorediate  species . 4 

3.  Podetia  ecorticate,  esorediate  (Subgenus  CLADINA) . 9 

4.  Podetia  yellowish,  usnic  acid  present 

(Subsection  UNCIALES) . 5 

4.  Podetia  grey-green  to  brownish,  usnic  acid  absent 

(Subsection  CHASMARIAE) . 7 

5.  Cartilaginous  cylinder  forming  an  unbroken  inner  lining  of  the 

podetia,  with  tiny  white  granules  resembling  pruina;  cortex  smooth 

and  shiny;  podetia  slender  (dry  habitats)  or  robust  (moist  habitats). 

Medulla  UV  +  blue-white  (squamatic  acid) . 126.  C.  uncialis 

5.  Cartilaginous  cylinder  more  or  less  discontinuous  or  fibrous;  cortex 
not  smooth  nor  shiny.  Medulla  UV-  (squamatic  acid  absent)  ...  .6 

6.  Podetia  inflated,  contorted,  perforate;  cartilaginous  cylinder  com¬ 
posed  of  loosely  interwoven  strands;  medullary  hyphae  (as  seen 
in  podetial  cross-sections)  loose  and  anastomosing,  (5-)6-8p,  in 

diameter . 124.  C.  boryi 

6.  Podetia  not  inflated  or  perforate;  cartilaginous  cylinder  composed 
of  closely  interwoven  strands;  medullary  hyphae  compact,  3-5 

(-7)  g.  in  diameter . 125.  C.  caroliniana 

7.  Soredia  present,  especially  at  podetial  tips;  podetia  usually  sparsely 

branched . 8 

7.  Soredia  absent;  podetia  intricately  branched . 122 .C.f areata 

8.  Soredia  usually  farinose,  scattered  in  irregular  patches  over  much 
of  the  podetium,  gradually  coalescing  into  a  continuous  sorediate 
area;  squamules  confined  to  the  lower  half  or  third  of  the  pode¬ 
tium,  or  entirely  absent . 121 .  C.  farinacea 

8.  Soredia  granular,  mostly  confined  to  the  tip  of  the  podetium; 

squamules  commonly  covering  entire  podetium.  Rare . 

. 120.  C.  scabriuscula 

9.  Branching  more  or  less  isotomic,  distinct  main  stems  absent  or  only 
exceptionally  present;  plants  giving  a  rounded,  tufted  appearance. 

Thallus  PD- . 10 

9.  Branching  anisotomic,  distinct  main  stems  usually  present . 11 

10.  Thallus  yellowish,  KOH-  (usnic  acid  present,  atranorin  absent); 
tetra-  to  pentachotomies  predominating,  usually  star  shaped 

around  an  axillary  hole . 128 .C.alpestris 

10.  Thallus  grey,  or,  rarely,  somewhat  yellowish,  KOH  +  yellow 
(usnic  acid  absent,  atranorin  present);  di-  or  trichotomies  pre¬ 
dominating,  axils  generally  closed.  Surface  appearing  very  rough, 


almost  tomentose;  algal  layer  not  continuous.  .  .  .  127.  C.  evansii 

11.  Thallus  PD- . 12 

11.  Thallus  PD  +  red  (fumarprotocetraric  acid) . 14 


12.  Thallus  KOH  +  yellow  (atranorin).  Thallus  surface  uneven, 


N>  N> 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


129 


appearing  “tomentose”,  algal  layer  very  discontinuous,  branch¬ 
ing  mostly  dichotomous  with  trichotomies  common.  Very  rare .  . 

. 130.  C.  terrae-novae 

12.  Thallus  KOH-  (atranorin  absent) . 13 

13.  Branches  very  robust,  often  sprawling;  axils  broadly  open;  branching 
usually  tetrachotomous  with  dichotomies  rare;  algal  layer  very  smooth 
and  compact,  appearing  almost  corticate.  Pseudonorangiformic  acid 

present.  Very  common . 133.  C.  submitis 

13.  Branches  usually  slender,  always  erect;  axils  often  closed  or  only 
slightly  open;  branching  usually  trichotomous  with  dichotomies 
common;  algal  layer  smooth  or  decomposed.  Pseudonorangiformic 

acid  absent.  Very  rare . 134 .C.mitis 

14.  Branching  predominantly  dichotomous,  tri-  and  tetrachotomies 
rare;  branchlets  usually  very  slender,  erect;  axils  infrequently 

open;  main  stems  often  indistinct;  pycnidial  jelly  red . 

. 129.  C.  subtenuis 

14.  Branching  predominantly  tri-  and  tetrachotomous  around  widely 
open  axils;  branchlets  robust,  falcate;  main  stem  always  distinct; 

pycnidial  jelly  colorless . 15 

15.  Thallus  blue-grey.  Usnic  acid  absent,  atranorin  present . 

. 131.  C.  rangiferina 

15.  Thallus  grey-green  to  yellowish  grey.  Usnic  acid  present,  atranorin 


absent.  . . 132.  C.  arbuscula 

16.  Podetia  and  apothecia  absent.  . . 17 

16.  Podetia  and/or  apothecia  present . 37 

17.  Medulla  PD  +  red,  orange,  or  yellow . 18 

17.  Medulla  PD-  . 30 

18.  Medulla  PD  +  red  ( fumarprotocetraric  acid) . 19 

18.  Medulla  PD  +  yellow  or  orange . 24 


19.  Squamules  sorediate  on  lower  surface,  broad,  entire  to  broadly  lobed 

. 20 


19.  Squamules  esorediate  on  lower  surface . 21 

20.  Squamules  large,  over  1 .0  mm  broad,  ascending . 

. 106.  C.  coniocraea 

20.  Squamules  minute,  0.5-0.75 (-1.0)  broad,  closely  appressed .  .  . 
. 49.  Lecidea  anthracophiia 


1.  Margins  of  squamules  finely  divided  to  ±  granulose . 22 

1 .  Margins  of  squamules  entire  to  broadly  crenate . 23 


22.  Grayanic  acid  (or,  very  rarely,  cryptochlorophaeic  acid)  present 

(in  Long  Island  material) . 103.  C.  chlorophaea 

22.  Grayanic  and  cryptochlorophaeic  acids  absent . 

. 112.  C.  caespiticia 

23.  Atranorin  present.  Squamules  strap-shaped;  margins  somewhat  revo¬ 
lute . 111.  C.  apodocarpa 

23.  Atranorin  absent . 

. C.  calycantha,  C.  clavulifera,  C.  mateocyatlia ,  C.  pyxidata 


<N  ri 


130 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


24.  Medulla  C  +  green . 93.  C.  strepsilis 

24.  Medulla  C- . 25 


5.  Medulla  KOH  +  blood  red  (norstictic  acid) . 96 .  C.  sabcariosa 

5.  Medulla  KOH-  or  KOH  +  yellow . 26 

26.  Medulla  KOH  +  deep  yellow  and  PD  +  orange  ( thamnolic  acid ) 

. 27 

26.  Medulla  KOH-,  PD  +  yellow . 29 

27.  Lower  surface  of  squamules  sorediate.  Terricolous,  corticolous,  or 

lignicolous . 83.  C.  macilenta 

27.  Lower  surface  of  squamules  esorediate.  Lignicolous . 28 

28.  Margins  of  squamules  granulose,  sometimes  reducing  the  primary 

thallus  to  a  granular  crust . 1 13.  C.  parasitica 

28.  Margins  of  squamules  finely  divided,  not  granulose . 

. 84.  C.  vulcanica 

29.  Squamules  entire  or  crenate.  Squamatic  and  baeomycic  acids  absent, 

psoromic  acid  present . 98.  C.  brevis 

29.  Squamules  finely  divided.  Squamatic  and  baeomycic  acids  present, 

psoromic  acid  absent . 1 16.  C.  atlantica  or  117.  C.  beaumontii 

30.  Thallus  C  +  red.  Lower  surface  of  squamules  sorediate . 

. 58.  Lee  idea  scalar  is 

30.  Thallus  C-  . 31 

31.  Upper  surface  or  lower  surface  of  squamules  yellow  or  yellowish 

(usnic  acid  present) . 32 

31.  Upper  surface  of  squamules  grey  to  grey-green,  lower  surface  white 

(usnic  acid  absent  or  not  detectable) . 34 

32.  Lower  surface  of  squamules  sorediate.  Squamatic  acid  present, 

barbatic  acid  absent . 86.  C.  incrassata 

32.  Lower  surface  of  squamules  esorediate.  Squamatic  acid  absent, 

barbatic  acid  present . 33 

33.  Squamules  very  large,  broadly  crenate  to  strap  shaped;  lower  surface 

yellowish.  Didymic  acid  absent . 92.  C.  robbinsii 

33.  Squamules  small,  usually  finely  divided;  lower  surface  white.  Didymic 

acid  present . 87.  C.  cristatella 

34.  Lower  surface  of  squamules  ±  sorediate.  Barbatic  acid  present, 

didymic  and  squamatic  acids  absent . 82.  C.  bacillaris 

34.  Lower  surface  of  squamules  esorediate . 35 

35.  Squamatic  acid  present,  didymic  acid  absent . 1 1 5.  C.  squamosa 

35.  Squamatic  acid  absent,  didymic  acid  present . 36 

36.  On  highly  decayed  wood  in  shaded  bogs . 85.  C.  didyma 

36.  On  soil,  dry  tree  bases,  or  dry  lignum  in  exposed  areas . 

. 87.  C.  cristatella 

37.  Apothecia  essentially  sessile  on  primary  squamules  or  on  very  short 
decorticate  podetia  (less  than  2  mm  tall);  squamules  finely  crenate; 
apothecia  brown,  flat  to  strongly  convex.  Squamules  PD  +  red 
(fumarprotocetraric  acid) . 112.  C.  caespiticia 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


131 


37.  Apothecia,  when  present,  on  ±  well  developed  podetia  at  least  parti¬ 
ally  corticate  and  over  2  mm  tall:  podetia  often  sterile . 38 

38.  Podetia  without  cups  or  tiers . 39 

38.  Podetia  with  more  or  less  distinct  cups  or  tiers . 64 

39.  Podetia  without  soredia  or  granules,  although  in  some  cases  somewhat 

ecorticate . 40 

39.  Podetia  granular  or  with  granular  or  farinose  soredia . 54 

40.  Apothecia  red;  podetia  and  squamules  yellowish  green  to  grey- 

green  (usnic  acid  present) . 41 

40.  Apothecia  brown,  tan,  or  huff,  or  absent;  podetia  and  squamules 
grey-green  to  olive-green  or  yellowish  green  (usnic  +  or— )  .  .  .42 

41.  Primary  squamules  covered  on  lower  surface  with  granular  or  farinose 

soredia.  Common  on  decaying  stumps  and  logs . 86.  C.  incrassata 

41.  Primary  squamules  without  soredia.  Very  common  on  many  sub¬ 
strates.  Podetia  usually  grey-green,  squamulose  on  bark  in  the  shade, 
and  yellow-green  without  squamules  on  the  ground  in  the  sun .... 

. 87.  C.  crist at ella 

42.  Podetia  more  or  less  abundantly  branched.  Podetia  PD  +  red  or 

orange . 43 

42.  Podetia  usually  simple,  or,  if  branched,  only  once  or  twice  near 

the  summit.  Podetia  PD  +  or  — . 44 

43.  Podetia  often  growing  in  dense  mats,  10-20  mm  tall,  often  bearing 
brown  apothecia;  holes  in  axils  often  surrounded  by  proliferations, 
giving  the  appearance  of  rudimentary  cups.  Podetia  KOH  +  lemon 

yellow,  PD  +  red-orange  or  orange-yellow  (thamnolic  acid) . 

. 1 18.  C.  f\oridana 

43.  Podetia  not  growing  in  dense  mats,  usually  over  20  mm  tall;  apothecia 
rare;  holes  in  axils  never  surrounded  by  proliferations.  Podetia  KOH- 


or  brownish,  PD  +  red  (fumarprotocetraric  acid)  .  .  .  122.  C.  furcata 

44.  Podetia  PD  + . 45 

44.  Podetia  PD  — . 52 

45.  Podetia  PD  +  red  (fumarprotocetraric  acid) . 46 

45.  Podetia  PD  +  yellow  to  orange . 48 


46.  Thallus  yellowish  green  to  grey-green;  podetia  7-15  mm  tall, 
minutely  squamulose;  apothecia  minute,  present  or  absent.  Very 

rare . 110.  C.  simulate i 

46.  Thallus  dark  or  pale  green-grey;  podetia  usually  less  than  10  mm 
tall,  not  squamulose;  apothecia  always  present,  large,  at  least 

equal  to  diameter  of  podetium.  Common . 47 

47.  Podetia  usually  grooved  and  twisted,  often  decorticate,  often  longi¬ 
tudinally  split  or  striate;  apothecia  buff  to  light  brown,  two  to  three 

times  the  diameter  of  the  podetium . 94.  C.  capitata 

47.  Podetia  usually  corticate,  verrucose  or  areolate,  not  twisted  or  striate; 
apothecia  dark  or  sometimes  light  brown,  one  to  two  times  the 
diameter  of  the  podetium . 97.  C.  clavulifera 


132 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


48.  Medulla  C  +  green,  KOH-  (strepsilin  and  baeomycic  acid). 

Thallus  and  podetia  olive  green;  podetia  ±  inflated . 

. 93.  C.  strepsilis 

48.  Medulla  C-,  KOH  +  or  —  (strepsilin  absent).  Thallus  and 
podetia  grey-green  or  brownish  green;  podetia  usually  slender.  . 

. 49 

49.  Podetia  with  perforate  tips  or  axils,  covered  with  large  or  small 

squamules  or  verrucae . 50 

49.  Podetia  not  perforate,  without  squamules,  or,  slightly  squamulose  on 

lower  half . 51 

50.  Podetia  KOH-  (baeomycic  acid  present).  Podetia  commonly  over 

10  mm  tall,  slender,  grey-green . 117.  C.  beaumontii 

50.  Podetia  KOH  +  yellow  (thamnolic  acid  present).  Podetia  usually 

under  10  mm  tall,  robust,  pale  grey  to  almost  white . 

. 1 14.  C.  santensis 


51.  Medulla  KOH  +  red  (norstictic  acid) . 96.  C.  subcariosa 

51.  Medulla  KOH—  (psoromic  acid) . 98.  C.  brevis 


52.  Thallus  with  a  distinct  yellow  tint  (usnic  acid  present) . 53 

52.  Thallus  without  any  hint  of  yellow  (usnic  acid  absent).  Thallus 
grey  or  brownish  green;  podetia  commonly  10-15  mm  tall,  fis¬ 
sured.  Atranorin  present.  Very  rare . 95.  C.  cariosa 

53.  Primary  squamules  small  (mostly  less  than  0.5mm  broad);  podetia 

common;  apothecia  flat,  reddish  brown,  abundant.  Rare . 

. 91.  C.  piedmontensis 

53.  Primary  squamules  very  large  (1-4  mm  broad);  podetia  rare,  very 
short,  arising  from  lateral  edges  of  squamules;  apothecia  strongly 

convex,  dark  brown.  Very  rare . 92.  C.  robbinsii 

54.  Podetia  PD  + . 55 

54.  Podetia  PD  - . 62 

55.  Podetia  PD  +  yellow  to  deep  yellow-orange,  KOH  +  lemon  yellow 

(thamnolic  acid) . 56 

55.  Podetia  PD  +  deep  red,  KOH—  or  +  dingy  brown  (fumarprotoce- 

traric  acid) . 58 

56.  Apothecia  brown  to  purple-brown,  common.  Podetia  and  margins 
of  primary  squamules  covered  with  large  corticate  granules .... 

. 1 1 3 .  C.  parasitica 

56.  Apothecia  red,  hut  sometimes  lacking . 57 

57.  Primary  squamules  esorediate;  podetial  soredia  coarsely  granular; 
podetia  often  decorticate  and  translucent  with  cartilaginous  layer 

exposed . 84.  C.  vulcanica 

57.  Primary  squamules  sorediate;  podetial  soredia  farinose  or,  rarely, 
granular,  covering  podetium;  podetia  often  decorticate  turning  brown 

to  black,  but  opaque . 83.  C.  macilenta 

58.  Podetia  short,  rarely  taller  than  6  mm,  with  blunt  apices,  covered 
with  coarsely  granular  soredia  on  the  lower  V%  to  %  of  podetium, 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


133 


and  farinose  soredia  on  the  upper  V2,  ecorticate  areas  abundant. 

Grayanic  acid  present . .  108.  C.  cylindrica 

58.  Podetia  usually  much  taller  than  6  mm,  apices  sharply  pointed, 
±  corticate  at  the  base,  corticate  on  upper  parts.  Grayanic  acid 

absent . 59 

59.  Podetia  partially  decorticate,  the  decorticate  areas  becoming  pellucid 
and  dark;  granular  soredia  covering  large  portions  of  the  podetia. 

Podetia  often  bent  or  contorted . 109.  C.  pityrea 

59.  Podetia  corticate  for  the  most  part,  or  the  cortex  is  replaced  by 

granular  or  farinose  soredia . . 60 

60.  Podetia  unbranched,  relatively  stout,  tapering  ±  abruptly  to  a 
sharp  point,  each  podetium  arising  from  the  center  of  a  primary 
squamule.  Podetia  and  squamules  with  a  vague  yellowish  green 
tint;  farinose  sorediate  on  upper  half  or  more  of  podetium; 
squamules  large,  sometimes  sorediate.  Common  and  variable .  . 

. 106.  C.  coniocraea 

60.  Podetia  commonly  branched,  long  and  slender,  not  arising  from 

the  center  of  primary  squamules . 61 

61.  Soredia  usually  farinose,  scattered  in  irregular  patches  over  much  of 
the  podetium,  gradually  coalescing  into  a  continuous  sorediate  area; 
squamules  confined  to  lower  half  or  third  of  the  podetium,  or  absent. 

Frequent . 121.  C.  farinacea 

61.  Soredia  granular,  mostly  confined  to  tip  of  podetium;  squamules  com¬ 
monly  covering  entire  podetium.  Rare . 120.  C.  scabriuscula 

62.  Podetia  corticate  for  most  of  length,  some  areas  bursting  into 
granular  soredia;  some  granular  soredia  on  lower  surface  of 
squamules  near  the  margins;  apothecia  red.  Usnic  acid  present 

or  absent.  Rare . 81.  C.  floerkeana 

62.  Podetia  mostly  sorediate,  often  with  many  decorticate  areas; 

apothecia  red.  Usnic  acid  absent . 63 

63.  Podetia  covered  with  granular  soredia,  or  soredia  becoming  farinose 
on  upper  half;  decorticate  areas  becoming  translucent,  then  brown; 

primary  squamules  esorediate.  On  wood  or  bark.  Rare . 

. . . 85.  C.  didyma 

63.  Podetia  entirely  covered  with  farinose  soredia,  occasionally  with  a 
small  corticate  area  at  the  base;  decorticate  areas  white,  opaque; 
primary  squamules  usually  having  granular  soredia  on  lower  surface 

near  the  margins.  On  various  substrates.  Very  common. . . 

. . . . . 82.  C.  bacillaris 

64.  Podetia  without  soredia  or  granules,  but  sometimes  squamulose 


or  minutely  verrucose . . . 65 

64.  Podetia  sorediate  or  granular. . .  .73 

65.  Cups  opening  into  podetia . 66 

65.  Cups  closed  by  continuous  membranes . 70 

66.  Podetia  KOH  +  deep  yellow  (thamnolic  acid!  . 67 


134  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

66.  Podetia  KOH- . 68 

67.  Cups  very  narrow,  almost  rudimentary,  slightly  perforated  at  tip. 

Rare . 114.  C.  santensis 

67.  Cups  broad,  with  extensive  proliferations . 123 .  C.  carassensis 

68.  Podetia  PD  + . 69 

68.  Podetia  PD  —  (squamatic  acid  present) . 115.  C.  squamosa 

69.  Podetia  PD  +  yellow  (baeomycic  and  squamatic  acids  present)  .  .  . 
. 1 1 6.  C.  atlantica 


69.  Podetia  PD  +  red  ( fumarprotocetraric  acid).  Podetia  and  cups  irreg¬ 


ularly  perforate  and  lacerate . 119.  C.  multiformis 

70.  Podetia  proliferating  from  center  or  edges  of  cups;  cups  shallow, 
flat,  or  slightly  convex;  podetia  corticate . 71 


70.  Podetia  simple,  deeply  goblet  shaped,  not  proliferating,  exten¬ 
sively  decorticate.  Inside  of  cup  lined  with  small  or  large  scattered 

areoles  or  flat  squamules.  Grayanic  acid  absent . 

. 102.  C.  pyxidata 

71.  Proliferations  irregular,  mostly  from  cup  edges;  cups  abortive,  ± 

squamulose;  squamules  large . 101.  C.  mateocyatha 

71.  Proliferations  from  center  of  cups,  regular;  cups  well  formed;  podetia 

esquamulose . . . 72 

72.  Cups  gradually  expanding  from  stalk;  podetia  usually  completely 

corticate.  On  neutral  soils . 99.  C.  verticillata 

72.  Cups  abruptly  expanding  from  stalk;  podetia  with  a  ±  continuous 
cortex  becoming  distinctly  areolate  or  partially  decorticate.  On 

acid  soils,  especially  in  or  near  bogs . 100.  C.  calycantha 

73.  Podetia  distinctly  yellowish  green  (usnic  acid  present  or  absent)  ...  74 

73.  Podetia  grey-green  or  brownish  (usnic  acid  absent).  Apothecia  brown 

. 77 

74.  Podetia  PD  +  orange  and  KOH  +  yellow  (thamnolic  acid) ;  usnic 
acid  absent.  Podetia  with  narrow,  shallow  cups,  corticate  at  base, 
soon  becoming  farinose  sorediate  and  sorediate  for  most  of 

length.  (On  Nantucket  Island,  not  on  Long  Island) . 

. [C.  digitata  (L.)  HofTm.] 

74.  Podetia  PD-,  KOH-;  usnic  acid  present . 75 

75.  Apothecia  brown.  Barbatic  acid  present,  zeorin  absent.  Cups  deep, 

goblet  shaped,  covered  with  farinose  soredia . 90.  C.  carneola 

75.  Apothecia  red.  Barbatic  acid  absent,  zeorin  present . 76 

76.  Cups  often  elongate,  somewhat  split  longitudinally;  soredia 

farinose.  Rare . 88.  C.  deformis 

76.  Cups  goblet  shaped,  not  split;  soredia  coarsely  granular.  Common 

. 89.  C.  pleurota 

77.  Soredia  coarsely  granular,  covering  entire  podetium.  Podetia  PD  + 
red  (fumarprotocetraric  acid)  or  PD-;  grayanic  or,  rarely,  crypto- 

chlorophaeic  acid  present . 103.  C.  chlorophaea 

77.  Soredia  farinose.  Podetia  PD  +  red  (fumarprotocetraric  acid)  .  .  .78 

78.  Cups  shallow,  deeply  dentate,  with  short,  spur-like  branchlets 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


135 


proliferating  from  edges  giving  a  star-like  appearance,  or,  infre¬ 
quently,  these  proliferations  are  lacking.  Homosekakaic  acid 
present  (but  often  difficult  to  demonstrate)  .  .  .  107.  C.  nemoxyna 
78.  Cups  usually  deep,  not  proliferating  from  edges.  Homosekakaic 

acid  absent . 79 

79.  Podetia  slender,  trumpet  shaped;  cups  narrow;  soredia  covering  entire 

podetium.  Substance  “H”  absent . 104.  C.  fimbriata 

79.  Podetia  broad,  goblet  shaped;  cups  wide;  soredia  usually  absent  on 
lower  half  of  podetium  where  there  is  a  continuous  cortex.  Substance 
“H”  present . 105.  C.conista 

36.  UMBiLICARIA 

1.  Thallus  pustulate;  undersurface  naked.  Apothecia  common;  disks  ± 
smooth,  becoming  somewhat  gyrose  with  age  with  margins  complete 

(leiodisc) . 137.  U.  papulosa 

1.  Thallus  smooth;  undersurface  rhizinate  or  lamellate . 2 

2.  Undersurface  with  flat,  reticulate  lamellae;  rhizines  absent;  apo¬ 
thecia  common;  disks  very  gyrose  with  margins  lacking  (actino- 

disc) . 136.  U.  muhlenbergii 

2.  Undersurface  densely  rhizinate,  with  a  mat  of  short  black 
rhizines;  apothecia  not  seen  on  L.  I.  material,  rare  elsewhere. 
(Disks  concentrically  gyrose  with  a  ±  complete  proper  margin 
[gyrodisc]) . 135.  U.  mammulata 

37.  SARCOGYNE 

1.  Apothecial  disks  rough,  verrucose,  carbonaceous;  epithecium  carbo¬ 
naceous,  thick,  very  uneven.  Apothecia  0. 3-1.0  mm  across;  hymenium 

(65-)  1 00- 1 20 ( -200 )  jj.  (including  the  black  epithecium) . 

. 140.  5.  simplex 

1.  Apothecial  disks  ±  smooth,  reddish  black  (especially  when  wet); 
epithecium  thin,  brown,  granular.  (Note:  Occasionally  some  carbo- 
aceous  material  appears  in  epithecium,  but  always  in  very  small 

amounts) . 2 

2.  Apothecia  0. 5-2.0  mm  across;  hymenium  85-1 20[j,  high;  hypo- 
thecium  usually  yellowish  or  brownish.  Common . 


. 138.  S.  clavus 

2.  Apothecia  less  than  1  mm  across;  hymenium  60-85 p  high;  hypo- 
thecium  hyaline.  Rare . 139.  S.  privigna 

39.  PERTUSARIA 

1.  Fruit  warts  smooth,  or  at  least  not  sorediate  or  granular . 2 

1 .  Fruit  warts  sorediate  or  granular . 8 

2.  Spores  8  per  ascus  (or,  rarely,  4  per  ascus) . 3 

2.  Spores  2  per  ascus  or  1  per  ascus . 5 

3.  Spores  uniseriate . 4 


3.  Spores  biseriate.  Fruit  warts  smooth;  ostioles  prominent,  depressed. 

Fruit  warts  PD  +  orange  and  KOH  +  red  (norstictic  acid) . 

. 145.  P.  propinqua 


136 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


4.  Thallus  epiphloedal,  thick;  fruit  warts  crowded,  over  1  mm  across, 

eupertusariate,  PD-,  KOH- . ..148.  P.  tuberculifera 

4.  Thallus  hypophloedal,  thin;  fruit  warts  scattered,  under  1  mm 

across,  ampliariate,  PD-,  KOH-  (?) . 142.  P.  alpina 

5.  Apothecium  lecanorine.  Disk  and  thallus  C  +  red.  .  .  .  149.  P.  velata 

5.  Apothecium  not  lecanorine;  fruit  warts  with  one  or  more  ostioles. 

Fruit  warts  and  thallus  C- . . . 6 

6.  Thallus  grey,  rarely  yellowish;  fruit  warts  eupertusariate,  poly- 

carpous  .  . . . . 7 

6.  Thallus  yellowish  green,  rarely  greyish;  fruit  warts  ampliariate 
(or,  infrequently,  somewhat  eupertusariate),  monocarpus  or 
dicarpous.  Thallus  UV  +  pink-orange;  fruit  warts  PD  ±  orange 

and  KOH  +  yellow  (stictic  acid) . 150.  P.  xanthodes 

7.  Fruit  warts  PD  +  red  ( fumarprotocetraric  acid).  Spores  (85-) 97- 

1 24 (-138)  x  35-45  [x,  always  hyaline,  radial  canals  absent . 

. . . 146.  P.  subpertusa 

7.  Fruit  warts  PD  +  orange  and  KOH  +  yellow  (stictic  acid).  Spores 
125-173  x  30-62  [x,  hyaline  to  brownish,  radial  canals  and  transverse 

wall  markings  usually  conspicuous . 151.  Melanana  macounii 

8.  Sorediate  warts  KC  +  violet.  Thallus  dark  ashy.  .  .  143.  P.  amara 

8.  Sorediate  warts  KC-.  Thallus  light  or  dark  grey.  .  . 9 

9.  Soredia  PD  +  orange  and  KOH  +  yellow  (thamnolic  acid) ;  spores 

2  per  ascus . . . 147.  P.  trachythallina 

9.  Soredia  PD-,  KOH-;  spores  1  per  ascus . .  10.  (P.  multipuncta) 

10.  Thallus  thin,  smooth;  fruit  warts  scattered,  bases  ±  broad;  spores 

97-1 10  x  45-48  [x . 144.  P.  cfr.  multipuncta  (#1 ) 

10.  Thallus  thick,  verrucose;  fruit  warts  crowded,  base  constricted; 
spores  125-150  x  (45-)55-70  |x .  .  .  .  144.  P.  cfr.  multipuncta  (#2) 

42.  IECANGRA 

1.  Thallus  becoming  distinctly  lobed  at  the  margins,  or  subfoliose. 

Saxicolous.  (Section  PLACODIUM) . . . 2 

1 .  Thallus  with  margins  not  lobed  or  subfoliose . .  3 

2.  Thallus  closely  adnate,  crustose;  apothecia  greenish  or  brownish, 

0.5-1. 5  mm  in  diameter.  On  calcareous  substrates . 

. . . 164.  L.  muralis 

2.  Thallus  ascending,  subfoliose  to  peltate;  apothecia  yellowish  or 
orange,  up  to  2.5  mm  in  diameter.  On  granite .  .  .  165.  L.  rubina 

3.  Apothecia  immersed  in  thallus  (especially  in  young  condition) ;  disks 
black;  spores  ( 1 2-)  16-20  x  7-lOjx.  Saxicolous  ( Section  ASPIC1LIA ) 

. . . . . . . 4 

3.  Apothecia  sessile  ( Section  LECANORA ) . . . . 5 

4.  Thallus  KOH-.  Pycnoconidia  (9-)  10-14  x  1  [x . 

. . . 154.  L.  caesiocinerea 

4.  Thallus  KOH  +  yellow  (stictic)  or  KOH  +  red  (norstictic) . 
Pycnoconidia  ( 10-)  13-18  x  1  ;x ............  .  .  157.  L.  cinerea 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


137 


5.  Disks  pitch  black.  Apothecia  up  to  2  mm  across;  epithecium  tinted 


violet,  especially  in  KOH;  spores  12-16  x  7-9  p, . 153.  L.  atra 

5.  Disks  yellowish  to  brown  or  dark  brown  (never  black) . 6 

6.  Spores  6-8  p,  wide . 7 

6.  Spores  2-6  (-7)  pi  wide . 11 


7.  Disks  heavily  pruinose,  C  +  orange;  apothecia  lavender.  Apothecial 

sections  KOH  +  blood  red  (norstictic  acid) . 

. 155.  L.  caesiorubella  subsp.  lathamii 

7.  Disks  epruinose  or  very  light  pruinose,  C-;  apothecia  brown.  Apo¬ 
thecial  sections  KOH  +  yellow  (atranorin  alone) . 8 

8.  Amphithecium  containing  large,  colorless  crystals;  epithecium 

inspersed  with  granules . 9 

8.  Amphithecium  without  large,  colorless  crystals;  epithecium  not 

inspersed  with  granules . 10 

9.  Epithecium  PD  +  red-orange  with  the  production  of  small  orange 
acicular  crystals.  Epithecial  granules  persistent  in  KOH;  apothecial 
disk  dark  brown,  epruinose,  strongly  convex;  margin  crenate,  soon 

becoming  thin  and  bead  like;  spores  12-14  x  7-8  pi.  Very  rare . 

. . . 160.  L.  degelii 

9.  Epithecium  PD  —  or  PD  +  yellow  (with  no  crystals  formed).  Epi¬ 
thecial  granules  dissolve  in  KOH;  apothecial  disk  yellow-brown  to 
red-brown,  often  slightly  pruinose,  flat  to  convex;  margin  thick, 

smooth  to  crenate;  spores  10-13  x  6-7  pi.  Very  common . 

. 156.  L.  chlarotera 

10.  Thallus  very  thick,  verrucose  and  chinky,  neither  granular  nor 
sorediate;  apothecia  up  to  2  mm  in  diameter,  often  twisted  and 
“urn  shaped”;  disk  reddish  brown;  apothecial  cortex  thick, 

45-50  pi.  On  cedar  stumps  and  old  wood . 163.  L.  laevis 

10.  Thallus  thinner,  smooth  to  granular  and  sorediate;  apothecia 
0.5-1. 0  mm,  circular,  closely  adnate;  disk  deep  mahogany  brown; 

apothecial  cortex  16-25  p,  thick.  On  bark . 169.  Lecanora  sp. 

11.  Spores  5-7  x  2-4  pi.  Apothecia  minute,  0. 2-0.4  mm  in  diameter.  On 

old  wood . 166.  L.  subintricata 

11.  Spores  8-16  x  3-7  p. . 12 

12.  Saxicolous  (on  limestone  and  mortar).  Thallus  almost  lacking; 
apothecia  0.25-0.50  mm  in  diameter;  disks  yellow-brown  to  olive- 
brown;  margins  white  or  ashy,  usually  persistent;  spores  9-10  x 

4-6  p, . 161.  L.  dispersa 

1 2.  On  bark,  wood,  or  bone . 13 

13.  Disks  yellow  pruinose,  lemon  yellow  when  young,  gradually  turning 
red-brown.  Thallus  well  developed,  grey,  very  rough;  spores  11-14  x 

4-5 p, . 159.  L.  cupressi 

13.  Disks  epruinose  or  lightly  white  pruinose,  yellow  to  brown . 14 

14.  Apothecial  margin  cortex  indistinct,  not  gelatinous;  thallus 
granulose  to  sorediate,  yellow-green;  apothecia  scattered  or 
crowded;  disks  yellow  to  buff . 16 


138 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


14.  Apothecial  margin  cortex  distinct,  gelatinous,  thick;  t  hall  us  essen¬ 
tially  absent,  or  if  present,  not  granular  or  sorediate;  apothecia 

very  crowded,  abundant . 15 

15.  Apothecial  sections  KOH-  (atranorin  absent).  Disks  buff  to  very  pale 

brown,  lightly  white  pruinose;  spores  10-13  x  3-5 ( -7 )  p. . 

.  162.  L.  hageni 

15.  Apothecial  sections  KOH  +  yellow  (atranorin  present).  Disks  yellow- 

brown  to  dark  brown,  epruinose;  spores  9-12  x  4-7  p. . 

. 168.  L.  cfr.  varia 

16.  Apothecial  margins  persistent,  becoming  thin  and  disappearing  in 
age,  soon  becoming  granulose;  spores  10-16  x  3-5  p..  Atranorin 

absent.  Frequent . 158.  L.  conizaea 

16.  Apothecial  margins  absent  in  all  but  the  youngest  apothecia, 
smooth  when  present  and  hardly  distinguishable  from  the  disk; 
spores  11-13  x  3-4  p..  Thallus  C-.  Very  rare..  .  .  167.  L.  symmicta 

43.  OCHROLECHIA 

1.  Thallus  (cortex  and  medulla)  C-  and  KC-.  Disks  often  somewhat 
pruinose;  spores  45-68  x  21-36  p..  Variolaric  acid  present  in  apothecial 


margin.  Common;  corticolous . 170.  O.  parella 

1.  Thallus  (cortex  or  medulla)  C  +  red . 2 


2.  Cortex  of  apothecial  margin  C  +  red;  amphithecial  medulla  C-. 
Thallus  thick,  verrucose;  spores  40-60  x  25-26  p..  Algae  present 
in  a  layer  (sometimes  not  continuous)  below  hypothecium; 

apothecial  cortex  relatively  thin.  Very  rare;  corticolous . 

. 171.  O.  rosella 

2.  Cortex  of  apothecial  margin  C-;  amphithecial  medulla  C  +  red, 
stipe  C-.  Thallus  thin,  rimose  to  verrucose;  spores  38-59  x 
21-26  p,.  Corticolous . 172.  Ochrolechia  sp. 

44.  HAEMATOMMA 

1.  Thallus  covered  with  sorediate  verrucae  towards  the  center,  becom¬ 
ing  smooth  at  the  edges;  thallus  eventually  becoming  a  granular 
sorediate  crust,  blue-grey  to  greenish  grey;  sterile.  Thallus  PD  + 
orange  and  KOH  +  yellow  (thamnolic  acid) . 174.  H.  sp. 

1 .  Thallus  coarsely  verrucose  or  almost  granular,  but  not  sorediate, 
whitish  green  to  yellowish  green;  apothecial  disks  red-brown,  com¬ 
mon;  spores  (35-)45-62  x  5-8  p..  Thallus  PD  +  orange  and  KOH  + 
yellow  (thamnolic  acid) . 173.  H.  ochrophaeum 

45.  CANDELARIELLA 

1.  Octosporous;  thallus  appearing  mostly  black,  or  pale  to  dull  yellow 
in  small  areas,  granulose  to  verrucose  or  subsquamulose.  On  cal¬ 
careous  rock . 175.  C.  aurella 

1.  Polysporous  (spores  about  20  per  ascus);  thallus  yo'k-  to  greenish 
yellow,  never  darkening,  granular-verrucose,  with  granules  or  sub¬ 
squamulose  verrucae  becoming  crowded  into  flattened  or  rounded 
patches.  On  granitic  rocks.  Often  sterile . 176.C.  vitellina 


139 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 

47.  PARMELIOPSIS 

1.  Thallus  yellow-green,  surface  bursting  into  irregular  laminal  soralia 
which  coalesce  into  a  mass  of  granular  soredia.  Thallus  PD-,  KOH-, 

usnic  and  divaricatic  acids  present . 179.  P.  ambigua 

1.  Thallus  grey-green  or  grey,  esorediate.  Thallus  PD  +  orange  and 
KOH  +  yellow  (thamnolic  acid);  usnic  and  divaricatic  acids  absent 

2 

2.  Thallus  isidiate,  adnate;  sterile . 178.  P.  aleurites 

2.  Thallus  not  isidiate,  often  ascending;  rarely  sterile . 

. 180.  P.  placorodia 


48.  PARMELIA 

1.  Thallus  yellowish  green  (usnic  acid  present) . 2 

1.  Thallus  greyish,  olive-green,  or  brownish,  no  trace  of  yellow  (usnic 

acid  absent) . 7 

2.  Soredia  in  punctiform  soralia,  or  tiny  verrucae  scattered  over 
upper  surface  of  thallus;  lobes  broad,  4-6  mm,  or  rarely  less. 
Medulla  PD  +  orange,  KOH-,  KC  +  red  ( protocetraric  and 

caperatic  acids  present).  Corticolous  or  saxicolous . 

. 184.  P.  caperata 

2.  Soredia  or  tiny  verrucae  absent;  lobes  less  than  4  mm  broad. 
Medulla  PD  +  yellow  or  orange,  KOH  +  red.  Saxicolous ....  3 


3.  Isidia  present.  Stictic  and  norstictic  acids  present . 4 

3.  Isidia  absent . 5 


4.  Lower  surface  of  thallus  black  almost  to  edge . 

.  185.  P.  conspersa 

4.  Lower  surface  of  thallus  buff  to  brown  throughout . 

. 193.  P.  plittii 

5.  Lower  surface  pale  brown  to  buff;  thallus  more  or  less  ascending. 

Salacinic  acid  present . 198.  P.  stenophylla 

5.  Lower  surface  black  except  very  close  to  margins . 6 

6.  Salacinic  acid  present . 202.  P.  tasmanica 

6.  Stictic  and  norstictic  acids  present . 182.  P.  arseneana 

7.  Thallus  olive-green  (wet)  or  brown  (dry),  never  grey.  Irregular 

laminal  soralia  present.  Medulla  C  +  red . 199.  P.  subaurifera 

7.  Thallus  grey  or  grey-green . 8 

8.  Pseudocyphellae  (white  dots)  scattered  over  upper  surface.  .  .  .9 

8.  Pseudocyphellae  absent . 12 

9.  Medulla  C-;  protolichesterinic  acid  present.  Soredia  or  isidia  absent, 

lower  surface  black,  becoming  pale  at  margins.  Very  rare . 

. 181.  P.  appalachensis 


9.  Medulla  C  +  red,  protolichesterinic  acid  absent . 10 

10.  Isidia  absent,  soredia  present . 11 

10.  Isidia  present,  soredia  absent.  Very  common.  ...  196.  P.  rudecta 


11.  Soredia  in  punctiform  soralia;  lower  surface  pale  brown.  Frequent 
.  200.  P.  subrudecta 


140 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


11.  Soredia  marginal;  lower  surface  black.  Very  rare . 

. 191 .  P.  olivetorum 

12.  Marginal  cilia  present . 13 

12.  Marginal  cilia  absent . 17 

13.  Soredia  absent . 14 

13.  Soredia  present . 15 

14.  Medulla  KC-,  KOH  +  red,  PD  +  yellow  (norstictic  acid  present, 
stictic  and  protocetraric  acids  absent);  cilia  usually  abundant; 
lower  surface  with  a  ±  broad  irregular  white  margin;  apothecia 

distinctly  perforate . 192.  P.  perforata 

14.  Medulla  KC  +  red,  KOH-,  PD  +  orange  (protocetraric  acid 
present,  norstictic  and  stictic  acids  absent);  cilia  very  sparse; 
lower  surface  of  thallus  uniformly  black,  lightening  to  brown  at 

margin;  apothecia  not  perforate . 190.  P.  michauxiana 

15.  Rhizines  present  to  the  thallus  edge  (hypotrachynoid) ;  upper  surface 
of  thallus  ±  covered  with  reticulate  cracks  and/or  tiny,  irregular 
white  areas  (maculae);  soredia  marginal  or  laminal.  Medulla  PD  + 

orange,  KOH  +  red  (salacinic  acid  present) . 195.  P.  reticulata 

15.  Rhizines  absent  from  edge  of  thallus  (amphigymnioid) ;  reticulate 

cracks  and  maculae  absent . 16 

1 6.  Lower  surface  of  thallus  smooth,  not  rugulose,  with  a  ±  broad, 
irregular  white  margin;  soredia  apical  or  marginal.  Medulla  KOH 
+  red,  PD  +  orange  (stictic  +  norstictic  acids).  Common.  .  .  . 

. 188.  P.  hypotropa 

16.  Lower  surface  of  thallus  rugulose,  uniformly  black  or  lightening 
slightly  to  brown  at  margin;  soredia  submarginal.  Medulla  KOH  + 
yellow,  PD  +  orange  (stictic  acid  present,  norstictic  acid  absent). 

Rare . 193.  P.  perlata 

Thallus  with  conspicuous  reticulate  ridges  and  depressions,  especi¬ 
ally  on  younger  portions  of  the  thallus.  Medulla  KOH  +  blood  red 

(salacinic  acid) . 18 

Thallus  ±  smooth,  rugose  or  cracked,  but  without  reticulate  ridges 

and  depressions.  Medulla  KOH  +  or  KOH  — . 19 

18.  Isidia  present . 197.  P.  saxatilis 

18.  Soredia  present  on  ridges . 201.  P.  sulcata 

19.  Soredia  present . 20 

19.  Soredia  absent . 22 

20.  Medulla  pale  yellow;  soredia  laminal.  Medulla  PD-  or  PD  + 

pale  yellow,  KOH  +  faintly  yellow . 183.  P.  aurulenta 

20.  Medulla  white:  soredia  marginal  or  laminal . 21 

21.  Medulla  PD  +  orange,  KOH-,  KC  +  red  (protocetraric  acid).  Sur¬ 

face  of  thallus  smooth  with  no  maculae;  lobes  mostly  3-4  mm  broad, 

crenate . 186.  P.  dilatata 

21.  Medulla  PD  +  orange,  KOH  +  red  (salacinic  acid).  Surface  of 

thallus  with  reticulate  cracks  and  maculae  (see  couplet  *15) . 

. 195.  P.  reticulata 


17 


17 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


141 


22.  Medulla  yellow,  especially  near  the  algal  layer;  chains  of  2  to  4 
moniliform  cells  scattered  throughout  the  medulla.  Medulla  PD 

+  orange,  KOH  +  orange  (unidentified  substance) . 

. 187.  P.  galbina 

22.  Medulla  white  throughout;  moniliform  cells  absent . 23 

23.  Medulla  PD  +  orange-red,  KOH-,  KC  +  red.  Rhizines  simple,  un¬ 
branched;  medulla  thick,  cottony  (see  couplet  #14) . 

. 190.  P.  michauxiana 

23.  Medulla  PD-,  KOH  +  red-brown.  Rhizines  branched;  medulla  ±  thin, 
not  cottony . . . 189.  P.  livida 

51.  CETRARSA 

1.  Terricolous.  Thallus  fruticose,  dark  brown,  with  broad  or  linear  lobes 
ascending  vertically,  producing  a  caespitose  growth  form;  pseudo- 

cyphellae  mostly  marginal,  linear.  Medulla  PD- . 

. . . . 207.  C.  islandica  subsp.  crispa 

1.  Corticolous.  Thallus  foliose,  brown,  grey,  or  yellowish;  lobes  often 

ascending  but  never  linear  and  never  caespitose . . 2 

2.  Thallus  grey,  pitted;  lower  surface  mostly  white,  sometimes 

mottled.  Medulla  I  +  blue . 208.  C.  tuckermanii 

2.  Thallus  yellowish  green  or  brownish  green,  never  grey . 3 

3.  Lower  surface  yellow;  upper  surface  greenish  yellow  .  .  .  209.  C.  viridis 

3.  Lower  surface  brown;  upper  surface  brown  or  greenish  brown.  .  .  .4 

4.  Apothecia  originating  on  upper  surface;  thallus  small,  appressed; 
lobes  narrow,  finely  divided,  0.5-0.75  mm  broad,  never  ciliate. 

Very  rare . . . . 206.  C.  fendleri 

4.  Apothecia  originating  on  lower  surface;  thallus  larger,  ±  ascend¬ 
ing;  lobes  1.5-4  mm  broad,  often  conspicuously  ciliate.  Common 

in  bogs.  Medulla  KC  +  red,  UV  +  (in  L.  I.  material) . 

. 205.  C.  ciliaris 

54.  ALECTORIA 

1 .  Thallus  caespitose,  wiry;  soralia  with  isidia.  Common . 

. 213  .A.  nidulifera 

1.  Thallus  pendent,  long;  soralia  without  isidia.  Very  rare . 

. . . 212  .A.  glabra 

55.  RAMALINA 

1.  Lacinae  subterete  or  angular,  ±  papillate.  Medulla  KOH  +  red  and 
PD  +  yellow  (salacinic  acid).  Spores  straight,  ellipsoid,  11-13  x  (4-) 

5-6  [j. . . . 217.  R.  willeyi 

1.  Lacinae  strongly  flattened . . . 2 

2.  Lacinae  with  elliptical,  delimited,  marginal  soralia  containing 
farinose  soredia.  Medulla  KOH-  or  KOH  +  red  (norstictic  acid 
[+ salacinic  acid  by  chromatography]).  On  roadside  trees  in 

Cape  Cod;  not  on  Long  Island . [R.  farinacea  (L.)  Ach.l 

2.  Lacinae  esorediate,  KOH-,  PD- .  . . 3 

3.  Lacinae  3-8  mm  broad,  coarsely  tuberculate-papillate.  Very  rare.  .  . 
. . . 214.  R.  cfr.  complanata 


142  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

3.  Lacinae  1-3  mm  broad,  not  tuberculate.  . . 4 

4.  Spores  fusiform,  straight  or  slightly  curved,  1 8-24 (-31)  x  3-5  p,; 
lacinae  strap  shaped,  with  white  striations  (pseudocyphellae?) 

usually  evident . 216.  R.  stenospora 

4.  Spores  ellipsoid,  straight,  8-13  x  4-6  p,;  lacinae  strap  shaped  to 
broadened,  often  with  numerous  short  proliferations  along  the 
margins;  smooth,  often  with  white  punctiform  pseudocyphellae, 
often  subcanaliculate . 215.  R.  fastigiata 


56.  USNEA 

1.  Medulla  rusty  red . 2 

1 .  Medulla  white . . . 3 

2.  Thallus  subpendent  to  pendent;  branching  irregular,  often  dichot¬ 
omous,  never  strigose;  isidiate-soralia  present;  apothecia  rare. 

Norstictic,  salacinic,  etc.,  absent . 219.  U.  mutabilis 

2.  Thallus  erect,  shrubby,  strigose;  branchlets  short;  isidia  and 
soredia  absent;  apothecia  common.  Norstictic  acid  present  in 

about  50  percent  of  the  specimens  seen . 220.  LI.  strigosa 

3.  Thallus  pendent,  filaments  exceedingly  slender,  never  tuberculate  or 

papillate;  stramineous  or  yellow-green.  Medulla  PD- . 4 

3.  Thallus  erect  or  subpendent;  filaments  generally  coarse,  papillae 
and/or  tuberculae  present;  dark  ashy  green,  at  least  in  older  portions. 

Medulla  PD  +  yellow  or  orange . 5 

4.  Branching  by  frequent  dichotomies;  perpendicular  side  branches 
infrequent;  axis  reddish  brown;  articulations  with  swollen  joints 
conspicuous;  cortex  intact.  Common  in  bogs.  .  .221.  U.  trichodea 

4.  Branching  infrequently  dichotomous;  perpendicular  side  branches 
common  and  regularly  spaced;  axis  white;  articulations  with 
swollen  joints  absent;  cortex  becoming  farinose.  Very  rare.  .  .  . 

. 218.  U.  longissima 

5.  Medulla  KOH-  (or  KOH  ±  very  faint  yellow).  Filaments  papillate 
and  tuberculate;  branches  coarse;  erect  or  subpendent;  cortex  very 
thick  and  chondroid;  isidiate-soralia  usually  present;  base  rarely 
blackened.  Protocetraric  acid,  or  rarely,  barbatic  or  fumarproto- 

cetraric10  acid  present .  222.  U.  sp. 

5.  Medulla  KOH  +  deep  yellow  or  red  (often  distinct  only  in  the 

apothecial  medulla) . 6 

6.  Isidiate-soralia  present;  medulla  very  lax;  apothecia  not  seen. 
KOH  +  red  or  yellow  (salacinic  or  stictic  acids  present).  Cape 

Cod  region,  fairly  common.  Not  on  L.  I . 

. [U.  cfr.  comosct  (Ach.)  Ach.] 

6.  Isidiate-soralia  absent;  medulla  compact;  apothecia  common. 
KOH  +  red  (norstictic  acid).  Filaments  strigose,  scrobiculate  on 
young  branches . 220.  U .  strigosa 

I0The  two  specimens  containing  fumarprotocetraric  acid  were  from  Cape  Cod 
(Brodo  4161.  4338). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  143 

57.  CALOPLACA 

1 .  Corticolous . 2 

1 .  Saxicolous . 6 


2.  Apothecial  margin  containing  few  or  no  algae;  thallus  yellow 
(KOH  +  red-purple),  thin,  sorediate.  Spores  13-17  x  8-10  p.; 

isthmi  5-7  p,  long . 227.  C.  discolor 

2.  Apothecial  margin  containing  a  distinct  algal  layer;  thallus 

esorediate . 3 

3.  Apothecial  disk  brown,  KOH-  (or  vaguely  pale  violet),  pruinose. 

Spores  13-19  x  7-lOp,;  isthmi  3-6  p,  long . 224.  C.  camptidia 

3.  Apothecial  disks  orange  or  yellow,  KOH  +  dark  purple  or  red-purple, 

not  pruinose . 4 

4.  Thallus  pale  yellow  or  cream  colored,  KOH  +  red-violet  (often 
weakly),  thin,  smooth.  Apothecial  disks  yolk  yellow  to  yellow- 
orange;  margins  yellow;  spores  11-13  x  4-6  p,;  isthmi  3-4  p,  long  .  . 

. 223.  C.  aurantiaca 

4.  Thallus  grey-green,  ashy,  or  dark  bluish  grey,  KOH- . 5 

5.  Amphithecium  thick,  ashy  to  blue-grey,  entirely  persistent;  apothecial 
disks  sordid  yellow  to  yellow-orange;  spores  12-16  x  7-8  p,;  isthmi 

(4-)5-6p.  long . 225.  C.  cerina 

5.  Amphithecium  very  thin,  pale  grey  to  ashy,  soon  disappearing  and 
revealing  an  orange  margin;  apothecial  disks  dark  orange  to  red- 
orange;  spores  10-14x4-7  p,;  isthmi  2-4 ( -5 )  p,  long.  .  .  230.  C.  pyracea 

6.  Spore  isthmi  less  than  3.5  p.  long;  thallus  minutely  areolate, 
yellow,  becoming  black  or  ashy,  disappearing.  Apothecia  0.25- 
0.40  mm  in  diameter;  disks  dark  orange  to  orange-brown;  margin 
yellow  to  orange,  often  becoming  leprose  or  granular;  spores 

12-17  x  7-9  p, . 228.  C.  feracissima 

6.  Spore  isthmi  more  than  3.5  p,  long;  thallus  yellow,  rarely  dark¬ 
ening  . 7 

7.  Thallus  effuse  granular  or  sorediate  to  subsquamulose  or  areolate. 
Apothecial  disks  orange;  margin  yellow,  often  sorediate;  spores  9-13 
x  5-7 (-9)  p,;  isthmi  3.5-5  p.  long;  sometimes  sterile.  .  .226.  C.  citrina 

7.  Thallus  smooth,  rimose,  areolate,  squamulose,  or  disappearing ....  8 

8.  Thallus  conspicuous,  squamulose;  apothecial  disks  dark  red- 
orange  to  orange-brown;  margins  dark  orange;  spores  11-17  x 

5-7  p, . 229.  C.  flavovirescens 

8.  Thallus  essentially  absent,  or  with  rare  yellow  squamules;  apo¬ 
thecial  disks  orange  with  yellow-orange  margins;  spores  11-15  x 
4-6  p. . 223.  C.  aurantiaca 


58.  XANTHORIA 

1.  Thallus  with  granular  soredia  in  labriform  soralia;  lobes  very  small 


and  narrow,  0.2-1. 0  mm  broad;  apothecia  rare . 23 1 .  X.  fallax 

1.  Thallus  esorediate;  lobes  broad,  (2-)3-4mm  broad,  flat;  apothecia 
common . 232.  X.  parietina 


144 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


59.  TELOSCHISTES 

1 .  Thallus  very  short,  caespitose,  yellowish  to  tan;  lacinae  flattened, 
±  striate,  giving  rise  to  short,  irregularly  shaped  side  branches  end¬ 
ing  in  pointed  cilia;  soredia  absent . 233.  T.  chrysophthalmus 

1.  Thallus  longer,  dark  yellow-orange;  lacinae  terete  or  ridged  and  angu¬ 
lar;  cilia  absent;  patches  of  soredia  frequent  throughout  length.  .  .  . 
. 234.  T.  flavicans 

60.  BUELLIA 

1 .  Saxicolous . 2 

1.  Corticolous  or  lignicolous . 3 

2.  Medulla  KOH  +  red  (norstictic  acid).  Thallus  whitish  grey, 
areolate;  prothallus  black,  well  developed.  Apothecia  mostly 

sessile  or  immersed  between  areoles . 239.  B.  stigmaea 

2.  Medulla  KOH-.  Thallus  dark  ashy  brown;  verrucose;  prothallus 

inconspicuous . 241.  B.  turgescens 

3.  Apothecial  sections  KOH  +  red  (norstictic  acid).  Apothecia  0.5- 

1.5  mm  in  diameter . 4 

3.  Apothecial  sections  KOH-.  Apothecia  less  than  0.5  mm  in  diameter 

. 5 

4.  Exciple  pallid  within;  grey  stipe  absent;  spores  17-24  x  6-8  [x; 

hymenium  80-130  [j.  high,  hyaline . 235.  B.  curtisii 

4.  Exciple  uniformly  dark;  grey,  T-shaped  stipe  present;  spores 
11-17  x  6-8  tx;  hymenium  55-75  [j.,  yellowish . 


. 240.  B.  stillingiana 

5.  Spores  8  per  ascus . 6 

5.  Spores  12  to  16  per  ascus.  Exciple  pale  within.  .  .  .237.  B.  polyspora 


6.  Thallus  PD  +  red  ( fumarprotocetraric  acid).  Spores  19-23  x 
8-9  ;x;  apothecial  margin  usually  absent;  disk  hemispherical.  .  . 

. 236.  B.  dialyta 

6.  Thallus  PD-.  Spores  9-11  x  6-7  |x;  apothecial  margin  distinct, 

disappearing  with  age;  disk  flat  to  slightly  convex . 

. 238.  B.  punctata 


61.  RINODINA 


1.  Saxicolous . 2 

1.  Corticolous . 4 

2.  Thallus  pale  grey  or  brownish  grey . 3 


2.  Thallus  yellowish  green.  Thallus  lobed  at  margins;  spores  10-12  x 

6-7  ix.  Medulla  C  +  red,  PD-,  KOH- . 245.  R.  oreina 

3.  On  siliceous  rock.  Thallus  verruculose  to  almost  squamulose;  spores 

17-23  x  9-13  [x . 243.  R.  confragosa 

3.  On  concrete.  Thallus  areolate  to  minute  verrucose;  spores  10-16  x 

6-8  [x . 247.  R.  salina 

4.  Spores  5-7 ( -8 )  x  (8-)  1 0- 1 2 ( - 1 5 )  |x;  hypothecium  dark  brown. 

Apothecia  less  than  0.5  mm  in  diameter . 244.  R.  milliaria 

4.  Spores  over  1 5  ix  long;  hypothecium  hyaline  or  yellowish . 5 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


145 


5.  Thallus  brownish  green  to  olive,  verrucose  or  granulose  to  smooth 
and  ±  squamulose;  spores  pachysporous  (examined  in  water),  16-23 

x  6-10  |x . 246.  R.  pachysperma 

5.  Thallus  thin,  smooth,  light  grey-green;  spores  mostly  pachysporous 

(examined  in  water),  17-24  x  9-11  jx . 242.  R.  applanata 

63.  PHYSCIA 

1 .  Thallus  deep  green,  olive-green,  or  brownish  grey;  upper  cortex  KOH- 

(atranorin  absent) . 2 

1.  Thallus  grey  or  grey-green;  upper  cortex  KOH  +  yellow  (atranorin 

present )  . 3 

2.  Thallus  esorediate;  lobes  finely  divided,  becoming  covered  with 
small  lobules;  lower  surface  white  to  buff;  rhizines  tan  to  brown; 

medulla  white.  Very  rare . 257.  Anaptychia  palmulata 

2.  Thallus  with  greenish  marginal  or  laminal  soralia;  lobes  crenate 
to  entire,  never  subsquamulose;  lower  surface  black;  rhizines 
black  with  white  tips;  medulla  white  (KOH-)  or  red-orange 

(KOH  +  purple).  Common . 252.  Physcia  orbicularis 

3.  Medulla  mustard  yellow.  Thallus  with  marginal  granular  soredia; 

lobes  pruinose,  1-2  mm  broad . 248.  Pyxine  sorediata 

3.  Medulla  white . 4 

4.  Thallus  sorediate  or  with  granules  resembling  soredia . 5 

4.  Thallus  esorediate.  Apothecial  disks  very  dark  brown  to  black, 

somewhat  pruinose . 10 

5.  Soredia  in  laminal  soralia . 255.  Physcia  tribacoides 

5.  Soredia  (or  granules)  marginal  or  terminal . 6 

6.  Lobes  helmet  shaped,  bursting  into  soredia.  Lobes  with  long, 

white,  marginal  cilia . 249.  Ph.  adscendens 

6.  Lobes  ±flat,  not  helmet  shaped . 7 

7.  Lobes  broad,  (2-)3-4mm,  rounded;  cortical  hyphae  parallel  to 

surface  .  8 

7.  Lobes  narrow,  0.3-2  mm  broad;  cortical  hyphae  at  right  angles  to 

surface  .  9 

8.  Lower  surface  white  (KOH-),  decorticate . 

. 258.  Anaptychia  pseudospeciosa 

8.  Lower  surface  yellow  (KOH  +  purple),  ecorticate . 

. . . 256.  Anaptychia  obscurata 

9.  Lobes  0.3-1 .0(-l .5)  mm  broad;  spores  16-19  x  6-9jx;  soredia  (or 
granules)  large,  marginal,  sometimes  reducing  thallus  to  a  granular 

crust.  Corticolous,  or  very  rarely,  saxicolous . 

. 251.  Physcia  millegrana 

9.  Lobes  very  narrow,  0.1-0. 5  mm  broad;  spores  12-16  x  6-8  [x;  soredia 
(or  granules),  marginal  and  apical,  occasionally  laminal.  Saxicolous 

. 254.  Ph.  subtilis 

10.  Medulla  KOH  +  yellow.  White  spots  (maculae)  present . 

. 250.  Ph.  aipolia 

10.  Medulla  KOH-.  White  spots  absent . 253.  Ph.  stellaris 


146 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


64.  ANAPTYCHIA 

1.  Thallus  esorediate,  brownish;  upper  cortex  KOH-  (atranorin  absent); 

lobes  finely  divided,  becoming  covered  with  small  lobules . 

. 257.  A.  palmulata 

1.  Thallus  sorediate,  greyish;  upper  cortex  KOH  +  yellow  (atranorin 

present);  lobes  not  finely  divided . 2 

2.  Lower  surface  light  to  deep  yellow,  KOH  +  red-violet,  not 

corticate . 256.  A.  obscurata 

2.  Lower  surface  white,  KOH-,  ±  corticate . 

. 258.  A.  pseudospeciosa 


65.  LEPRARIA 

1.  Saxicolous.  Thallus  grey  to  dark  ashy  green;  granules  large,  often 
forming  a  ±  lobed,  zonate  thallus.  Thallus  PD  +  red,  and  KOH- 
(fumarprotocetraric  acid),  or  rarely,  PD  +  yellow  (barbatolic  acid) 

. 260.  L.  zonata 

1.  Corticolous  or  lignicolous . 2 

2.  Thallus  with  a  distinct  bluish  grey  cast,  a  ±  thin  layer  of  dis¬ 
persed  granules  with  little  or  no  prothallus.  Thallus  KOH  + 
yellow  and  PD-  (atranorin)  or  rarely,  KOH  +  yellow  and  PD 
+  red  (fumarprotocetraric  acid  +  atranorin)  .  .  .  .259.  L.  incana 
2.  Thallus  pale  green  or  sometimes  yellowish  green,  thick  masses  of 
granules  subtended  by  a  thick,  white,  prothalline  mat.  Thallus 
KOH  +  yellow  and  PD  +  orange  (atranorin  +  stictic  acid)  .  . 
. 261.  L.  sp. 


ARTHOPYRENIACEAE 

1.  ARTHOPYRENIA  Mass. 

1.  Arthopyrenia  cerasi  (Schrad.)  Mass.  Ricerch.  Auton.  Lich.  167. 
1852.  Verrucaria  cerasi  Schrad.  Ann.  d.  Bot.  22:86.  1797. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2375  (123). 

Fink  (1935)  reports  A.  cerasi  from  young  oaks,  but  the  Long  Island 
material  was  on  Myrica  pensylvanica. 

Distribution  —  Maryland,  Iowa,  California  (Fink,  1935);  Europe. 

2.  Arthopyrenia  pinicola  (Hepp)  Mass.  Symm.  Lich.  118.  1855. 
Pyrenula  punctiformis  var.  cineropruinose  f.  pinicola  Hepp,  Flecht.  Europ. 
106.  1853. 

Material  seen  —  SUFFOLK  COUNTY :  Brodo  3176  ( 65 ) . 

Degelius  ( 1941 )  discusses  the  synonomy  and  gives  a  detailed  descrip¬ 
tion  of  his  specimens.  The  Long  Island  material  fits  his  description  very 
closely.  It  was  collected  on  the  base  of  a  white  oak  ( Quercus  alba). 
I  also  found  a  specimen  on  Uhnns  americana  in  central  New  York  State 
( Brodo  VIII,  in  herb.  CAN). 

Distribution  —  Tennessee;  Europe. 

NOTE:  Arthopyrenia  halodytes  (Nyl.)  Arn.  (A.  sublitoralis  [Leight.] 
Arn.)  not  collected,  but  may  be  found  on  shells  and  barnacles  if  sought. 


147 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 

2.  LEPTORHAPHIS  Korb 

3.  Leptorhaphis  epidermidis  (Ach.)  Th.  Fr.  Nova  Acta  Reg.  Soc.  Sci. 
Upsal.  III.  3:373.  1861.  (Lich.  Arct.  273.  1860.)  Lichen  epidermidis 
Ach.  Lich.  Suec.  Prodr.  16.  1798. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1015  (27),  1120 
(78),  1395  (65),  1985  (91),  2455  (22),  2591  (97),  2773  (31),  3100 
(122),  3817  (66). 

All  specimens  of  this  species  were  found  on  the  bark  of  Betula 
populifolia.  It  is  similar  to  Polyblastiopsis  quercicola  in  gross  morpho¬ 
logical  features  such  as  shape,  size,  and  position  of  the  perithecia,  but 
their  spores  and  substrates  are  quite  different. 

Distribution  —  Eastern  United  States  (Fink,  1935);  Connecticut, 
Michigan,  Wisconsin,  Arizona,  Black  Hills:  Temperate  element,  East 
Temperate  subelement;  Europe. 

3.  POLYBLASTIOPSIS  Zahlbr. 

4.  Polyblastiopsis  quercicola  sp.  nov. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2651  (61),  2674 
(108),  2788  (31). 

Thallus  subtilissimus,  hypophloedalis,  albus;  algae  non  visa. 
Pseudothecia  nigra,  diam.  0.15-0.25  mm,  hemispherica,  dispersa, 
superficialia,  sed  saepe  ex  parte  thallo  vel  epidermide  corticis  tecta; 
ostioles  conspicues  saepe;  parietes  carbonisati,  virides-nigres  lit 
oblinentur,  praecipuus  in  KOH.  Filamentae  paraphysoideae  et  asci 
immersi  in  substrato  gelatinoso,  dissoluto  in  KOH.  Filamentae  para¬ 
physoideae  persistentes,  distinctae  in  KOH,  ramosa  et  anastomosa 
copiose,  1.5-2. 5  g.  diam.  Asci  parietes  ±  snbtiles  in  H,0.  sed  perspicue 
crassi  in  KOH ,  praecipuus  apice.  Sporae  octonae,  irregulariter  seri- 
atae,  morales  vel  submurales,  septis  transversis  3-6,  septis  longitudi- 
nalibus  1-2,  hyalinibus;  vagina  episporis  lucida,  conspicua,  levis  in 
HjO,  ±  irregularis  in  KOH;  16-27  a  7-10  g.  Pycnidia  nigra,  minu- 
tissima,  dispersa.  Pycnoconidia  hyalina,  non  septata,  elongata- 
cylindrica  ad  fusiforma,  7-9  x  1  g.  Ad  corticem  Quercus  alba. 
Holotype:  New  York.  Suffolk  County:  Shoreham.  Saint  loseph's 
Villa,  N.  Country  Road,  black  oak  woods,  Brodo  2651,  July  7,  1961,  on 
Quercus  alba,  0  ft.  and  higher  (MSC)  (see  figures  82,  83,  87d). 

Thallus  very  thin,  hypophloedal,  appearing  white;  no  algae  evident. 
Pseudothecia  black,  0.15-0.25  mm  in  diameter,  hemispherical,  scattered, 
superficial,  but  often  partially  covered  by  thallus  and/or  upper  layers  of 
bark;  ostioles  often  conspicuous;  walls  carbonaceous,  greenish  black  when 
smeared,  especially  in  KOH.  Paraphysiod  threads  and  asci  embedded  in 
gelatinous  material  which  dissolves  in  KOH;  paraphysoid  threads  persist¬ 
ent,  distinct  in  KOH,  abundantly  branched  and  anastomosing,  1 .5-2.5  g 
in  diameter.  Asci  appearing  ±  thin  walled  when  mounted  in  water,  but 
clearly  thick  walled,  especially  at  apex,  when  mounted  in  KOH.  Spores 
8  per  ascus,  irregularly  arranged,  muriform  or  submuriform,  3  to  6  trans- 


148 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


verse  septa,  1  to  2  longitudinal  septa,  hyaline;  hyaline  epispore  sheath 
conspicuous,  smooth  in  a  water  mount,  ±  irregular  in  a  KOH  mount; 
16-27  x  7-10 1 jl .  Pycnidia  black,  extremely  minute,  scattered.  Pycnoco- 
nidia  hyaline,  nonseptate,  elongate-cylindrical  to  fusiform,  7-9  x  1  [j.. 

The  specimens  from  Long  Island,  as  well  as  some  from  New  Jersey 
(Brodo  3728,  3755),  were  rather  uniform  in  morphology.  The  thallus 
often  covers  several  square  centimeters,  or  even  decimeters,  on  or  near 
the  bases  of  oaks.  Pseudothecia  vary  little  in  size.  Spores  are  16-27  x 
7-10  [j.  and  always  show  the  gelatinous  epispore  sheath.  Pycnoconidia  are 
6-9  x  1 .0-1.5  (A,  hyaline  and  nonseptate.  Algae  (apparently  Trentepohlia ) 
were  in  very  small  amounts  just  below  the  pseudothecia  of  a  few 
specimens. 

The  species  differs  from  similar  P.  fallaciosa  (Stizenb.)  Zahlbr.  in 
having  larger  spores  and  hyaline,  nonseptate  rather  than  brown,  septate 
pycnoconidia,  and  from  P.  lactea  (Mass.)  Zahlbr.  in  having  somewhat 
smaller  spores  and  eight  rather  than  four  spores  per  ascus.  Polyblastiopsis 
fallax  (Nyl.)  Fink,  which  is  close  to  P.  quercicola  from  the  description  in 
Fink  (  1935),  appears  to  be  a  synonym  of  Arthopyrenia  fallax  (Nyl.)  Arn. 

The  species  was  found  on  the  bark  of  Quercus  alba  and  Q.  stellata. 
Similar  species  are  almost  always  found  on  Betula  or  some  other  tree,  and 
so  the  unusual  substrate  served  as  the  source  of  the  specific  epithet. 

Distribution  —  New  Jersey  (see  above). 


Figure  82.  Polyblastiopsis  quercicola  (holotype).  Scale  equals  1  mm. 
Drawing  by  Brenda  Carter  Haas. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


149 


Figure  83.  Polyblastiopsis  quercicola  (holotype).  (a)  ascospore,  (b) 
paraphysoid  threads,  (c)  developing  and  mature  asci.  Scales  equal 
10  p,.  Drawings  by  G.  Morgen-Jones,  with  the  aid  of  a  camera  lucida 
apparatus,  from  material  mounted  in  water. 


150 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


ARTHONIACEAE 

4.  ARTHONIA  Ach. 

5.  Arthonia  caesia  (Flot.)  Korb.  Parerg.  Lich.  269.  1861.  Conian- 
gium  caesium  Flot.  in  Korb.  Syst.  Lich.  Germ.  295.  1855. 

Material  seen  —  SUFFOLK  COUNTY:  19  specimens  collected  by 
Imshaug  and/or  Brodo. 

Fink  (1935)  probably  included  this  species  in  his  concept  of 
A.  impolita  (Ehrh.)  Borr.  (Syn.  A.  pruinosa  Ach.).  The  two  species  are 
similar  in  their  leprose  thalli  and  small,  dark,  heavily  blue-grey  pruinose 
ascocarps,  but  may  be  separated  as  follows:  A.  impolita :n  phycobiont 
Trentepohlia,  fruit  base  hyaline,  thallus  reactions  KOH  yellow  and  KC 
red;  A.  caesia:  phycobiont  Trebouxia,  fruit  base  yellow  to  red-brown, 
thallus  reactions  negative  with  KOH  and  KC. 

The  Long  Island  material  was  mostly  fertile  and  agreed  in  all 
respects  with  European  descriptions  of  the  species. 

The  species  is  found  on  bark  of  all  kinds  in  shaded  or  exposed 
woods,  or  on  exposed  downs.  Under  the  right  conditions,  it  apparently 
has  a  very  rapid  growth  rate  and  was  seen  almost  covering  young  twigs 
and  branches  in  an  oak  forest  in  Laurel. 

Distribution  ■ —  Tennessee,  Wisconsin,  but  probably  common  in  east¬ 
ern  United  States;  Europe. 

6.  Arthonia  mediella  Nyl.  Not.  Soc.  Faun.  FI.  Fenn.  Forhandl.  1:238. 
1858-59. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  795 A  (90B). 

The  Long  Island  material  agreed  well  with  the  description  by  Redin¬ 
ger  ( 1937-38),  and  with  a  specimen  from  Finland  ( Lang  347 ,  hb.  MICH). 

Redinger  states  that  the  species  is  corticolous,  but  the  Long  Island 
specimen  was  on  old  wood  on  an  exposed  bluff  overlooking  Long  Island 
Sound. 

Distribution  —  First  North  American  record;  Europe. 

7.  Arthonia  polymorpha  Ach.  Syn.  Lich.:  7.  1814. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1070  (98). 

This  species  is  similar  to  A.  siderea  in  the  color  and  stellate  arrange¬ 
ment  of  the  hysterothecia.  It  is,  however,  readily  distinguished  by  its 
pruinose  hysterothecia  and  pale  brown  spores  with  equal  sized  cells. 

It  was  found  on  the  bark  of  Carya  glabra. 

Distribution  —  Maryland,  Florida,  Louisiana,  Illinois,  Iowa,  and 
California  (Fink,  1935). 

8.  Arthonia  punctiformis  Ach.  Kgl.  Vet.  Akad.  Nya  Handl.  130.  1808. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25742  ( 132). 

Orient,  Latham  782.  April  5,  1914,  (Latham);  Montauk,  Latham  3953, 
April  6,  1927  (Latham);  Greenport,  Latham  8608,  April  30,  1939 
(Latham);  Orient,  Latham  24178,  March  21,  1915  (Latham). 


11  Based  on  Michigan  material  (MSC). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


151 


This  species  is  similar  to  A.  radiata  (Pers.)  Ach.  in  many  respects, 
but  the  latter  has  larger  ascocarps  and  an  epiphloedal  thallus,  whereas 
the  ascocarps  of  A.  punctiformis  are  0.1-0. 2  mm  across  and  its  thallus 
is  hypophloedal. 

Arthonia  punctiformis  is  found  on  hark  of  various  kinds,  and  is  rare 
on  Long  Island. 

Distribution  —  Maine,  Connecticut,  Tennessee,  Minnesota,  Alaska; 
throughout  the  United  States  (Fink,  1935);  Europe;  Asia  (Vainio,  1928). 

9.  Arthonia  sexlocularis  Zahlbr.  Ann.  Myc.  12:  336.  1914. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2818  (115). 

This  single  specimen  was  found  growing  on  the  deeply-shaded  base 
of  Celtis  sp.  The  distinctive  spores  agree  perfectly  with  the  description 
of  Redinger  (1937-38)  based  on  European  material. 

Distribution  —  First  North  American  record;  Europe. 

10.  Arthonia  siderea  Degel.  Ark.  Bot.  30A(1):  14.  1940. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1182  (101),  1203 

(101),  1515  (100B),  1862  (117),  1874  (117),  2306  (93),  2419  (113), 
2606  (84),  2626  (71),  2686  (110),  2727  (111),  3247  (119);  Orient; 
Latham  8598A,  April  10,  1939  (Latham);  Orient,  Latham  8600,  April 
30,  1939  (Latham). 

Although  Degelius  described  this  species  as  having  “black  apo- 
thecia,”  the  hysterothecia  actuaVy  range  in  color  from  red-brown  to  black. 
When  moistened,  the  hysterothecia  always  appear  a  deep  mahogany  and 
never  are  black.  The  holotype,  which  Dr.  Degelius  kindly  sent  me,  had 
hysterothecia  which,  although  almost  black  when  dry,  also  show  the  red- 
brown  tint  when  wet.  The  Long  Island  material  agrees  in  every  respect 
with  the  holotype. 

This  species  is  probably  more  common  than  would  be  thought, 
judging  from  the  number  of  times  it  has  been  reported.  In  Fink  (1935), 
A.  siderea  easily  keys  out  to  A.  gregaria  (Weig.)  Korb.  which  is  now 
recognized  as  a  synonym  of  A.  cinnabarina  (DC.)  Wallr.  (see  discussion 
in  Redinger,  1937-38).  Arthonia  cinnabarina,  however,  has  rusty  red, 
often  powdery  hysterothecia,  which  are  KOH  +  red-violet.  The  hystero¬ 
thecia  of  A.  siderea  are  KOH-,  smooth  and  often  shiny.  The  spore  type 
and  ascocarp  color  clearly  distinguish  it  from  A.  radiata  (Pers.)  Ach. 

Arthonia  siderea  is  found  on  the  bark  of  black  oaks  well  above  the 
base,  and  is  probably  photophilous  (figure  67). 

Distribution  —  Maine;  endemic. 

5.  ARTHOTHEL1UM  Mass. 

11.  Arthothelium  taediosum  (Nyl.)  Miill.  Arg.  Flora  63:  287.  1880. 
Arthonia  taediosa  Nyl.  Ann.  Sci.  Nat.  Bot.  IV.  3:  171.  1855. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3832  (66);  Green- 
port,  Latham  8610,  April  16,  1938  (Latham);  Greenport,  Latham  7206, 
January  26,  1933  (MICH). 

Two  of  the  specimens  from  Long  Island  differ  somewhat  in  spore 
size  (32-37  x  10-17  [a  in  Brodo  3832;  34-46(-55)  x  14-23  (-27)  p,  in 


152  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Latham  8610),  but  the  thallus  and  ascocarp  characters  agree  perfectly. 
Except  for  the  large  spore  sizes  in  the  Latham  specimen,  the  specimens 
fit  the  description  of  A.  taediosum  given  by  Redinger  (1937-38)  very 
well.  Redinger’s  spore  measurements  are  24-30(-33)  x  8-13  t j. .  The  only 
other  possible  Arthothelium  which  could  be  considered  for  the  Latham 
specimen  would  be  A.  distendens  (Nyl.)  Mull.  Arg.,  and  Nearing  anno¬ 
tated  the  specimen  with  that  name.  After  having  examined  a  number 
of  specimens  of  both  species  in  the  Fink  herbarium  (MICH)  I  feel 
there  is  little  doubt  that  both  Long  Island  specimens  belong  to  A.  taedio¬ 
sum.  I  could  find  no  spores  in  Latham  7206,  but  the  specimen  was  iden¬ 
tified  by  Josiah  Lowe  as  this  species.  The  specimens  of  A.  distendens  had 
thicker  thalli  and  much  broader  hysterothecia  (up  to  1  mm  across).  In 
the  Long  Island  material,  the  ascocarps  were  punctiform  to  irregular,  and 
0.1 -0.2  mm  across. 

Arthothelium  taediosum  was  found  on  the  smooth  bark  of  red  maple 
and  oak  on  Long  Island. 

Distribution  —  Eastern  United  States,  California  (Fink,  1935); 
Connecticut;  Europe  (Redinger,  1937-38). 

6.  MICAREA  Fr. 

12.  Micarea  melaena  (Nyl.)  Hedl.  Bih.  Kgl.  Svensk.  Vet.  Akad. 
Handl.  afd.  Ill,  no.  3,  18:  82.  1892.  Lecidea  melaena  Nyl.  Bot.  Not.  182. 
1853. 

Material  seen  —  NASSAU  COUNTY:  Brodo  1510  (14). 

This  species  is  considered  in  the  genus  Bilimbia  in  Fink  (1935). 

The  Long  Island  specimen  was  found  on  a  rotting  stump. 
Distribution  —  Eastern  United  States  (Fink,  1935);  Connecticut, 
Michigan;  Temperate  element,  East  Temperate  subelement;  Europe;  Asia 
(Vainio,  1928). 

13.  Micarea  prasina  (Fr.)  Korb.  Syst.  Lich.  Germ.  399.  1855. 
Biatora  prasina  Fr.  Stirp.  Agri.  Ferns.  38.  1826. 

var.  sordidescens  (Nyl.)  comb.  nov. 

Lecidea  sordidescens  Nyl.  Flora  57:  312.  1874. 

Material  seen  —  NASSAU  COUNTY:  Brodo  1497  (9),  3497  (4). 
SUFFOLK  COUNTY:  Brodo  7959(51),  2578  (13). 

The  Long  Island  material  would  fall  into  the  var.  sordidescens  (Nyl.) 
Lettau  emend.  Erichs,  of  Catillaria  prasina  (Fr.)  Th.  Fr.  (Erichsen,  1957). 
This  variety  has  a  KOH  +  violet  “epithecial”  reaction.  The  spores  are 
either  mostly  nonseptate  or  mostly  uniseptate,  but  both  types  can  always  be 
found  in  a  smear  of  the  ascocarp.  The  species  is  treated  in  the  genus 
Catillaria  in  Fink  (1935). 

Micarea  prasina  is  found  on  rotten  wood,  but  is  rare  on  Long  Island. 
Distribution  - —  Northern  and  eastern  United  States  (Fink,  1935); 
Connecticut,  Minnesota,  Black  Hills:  Temperate  element,  North  Tem¬ 
perate  subelement;  Europe;  Asia  (Vainio,  1928). 


153 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 

OPEGRAPHACEAE 

OPEGRAPHA  Ach. 

14.  Opegrapha  cinerea  Chev.  Journ.  Phys.  Chim.  Hist.  Nat.  94:  41. 

1822. 

Material  seen  — SUFFOLK  COUNTY:  Imshaug  25674  (72),  Brodo 
59-221  (72),  785  (90A),  797  (90B),  1079  (98),  1755  (127),  2650  (61); 
Greenport,  (collector  unknown),  April  1903  (FH). 

This  species  is  somewhat  similar  to  O.  vulgata  (Ach.)  Ach.,  which  is 
distinguished  by  having  (1)  unbranched,  shiny  hysterothecia,  (2)  a  green¬ 
ish  brown  thallus,  (3)  spores  15-20;j.  long,  and  (4)  a  specificity  for 
coniferous  bark  (Redinger,  1937-38). 

Opegrapha  cinerea  is  found  on  the  bark  of  smooth-barked  broadleaf 
trees  such  as  Carya  sp.  and  Quercus  sp.  (which  agrees  with  the  habitat 
notes  of  Redinger,  1937-38).  All  the  Long  Island  specimens  were  found 
within  a  half  mile  of  the  north  shore  (figure  62). 

Distribution  —  Florida  (Fink,  1935),  the  Smoky  Mountains  of 
Tennessee;  Europe. 

15.  Opegrapha  rufescens  Pers.  Neue  Ann.  Bot.  1:29.  1794. 
Material  seen  —  SUFFOLK  COUNTY:  Orient  Point,  Latham, 

June  5,  1911  (NYS). 

Distribution  —  Florida  (Fink,  1935);  Europe. 

CALICIACEAE 

8.  CHAENOTHECA  (Th.  Fr.)  Th.  Fr. 

16.  Chaenotheca  phaeocephala  (Turn.)  Th.  Fr.  Nova  Acta  Reg.  Soc. 
Sci.  Upsal.  Ill,  3:  351.  1861.  (Lich.  Arct.  251.  1860).  Lichen  phaeo- 
cephalus  Turn.  Trans.  Linn.  Soc.  Lond.  8:  260.  1807. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25810  (86),  Brodo 
2124  (102). 

The  spores  of  this  species,  normally  brown,  often  become  colorless 
in  KOH. 

Chaenotheca  phaeocephala  is  rare  on  Long  Island  and  is  restricted 
to  rotting  stumps  of  Chamaecyparis  thyoides  in  shaded  bogs.  It  was  col¬ 
lected  once  in  southern  New  Jersey  (Brodo  3772)  and  once  on  Cape  Cod 
( Brodo  4337 )  in  similar  habitats  and  on  the  same  substrate. 

Distribution  —  New  England  and  Minnesota  (Fink,  1935);  Michi¬ 
gan:  Temperate  element,  North  Temperate  subelement(?) ;  Europe. 

VERRUCARIACEAE 

9.  VERRUCARIA  Schrad. 

17.  Verrucaria  microspora  Nyl.  Ann.  Sci.  Nat.  Bot.  IV.  3:  175.  1855. 
Verrucaria  subsuperficialis  Fink  in  Hedr.  Mycologia  25:  304,  1933. 

Material  seen  —  SUFFOLK  COUNTY:  Orient,  Latham,  1925 
(holotype  of  V.  subsuperficialis)  (MICH);  Orient,  Latham,  1927, 

(MICH,  FH). 


154  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  holotype  of  V.  subsuperficialis  was  compared  with  a  specimen 
of  V.  microspora  from  Denmark,  which  was  kindly  sent  to  me  by  Dr. 
Degelius.  The  two  specimens  were  identical  in  morphology  and  ecology, 
both  having  thin,  membranous,  dark  brown  thalli,  small  spores,  and  both 
having  been  found  in  the  hydrohaline  stratum  (p.  61)  on  quartz  pebbles. 

Distribution  —  Maine:  Temperate  element,  Maritime  subelement; 
maritime  Europe  (Santesson,  1939;  des  Abbayes,  1934). 

18.  Verrucaria  muraiis  Ach.  Meth.  Lich.  115.  1803. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2833  (115). 

This  species  was  found  growing  alongside  V.  nigrescens  on  mortar 
and  brick  in  the  aerohaline  stratum  at  Orient  Point.  It  differs  from  the 
latter  species  in  having  a  white  or  ashy  thallus,  larger  spores,  and  an 
entirely  different  type  of  perithecium.  Its  involucrellum  is  black,  hemi¬ 
spherical,  and  is  almost  entirely  external  to  the  thallus;  the  exciple 
appears  hyaline.  The  species  fits  the  description  in  Zschacke  (1933) 
fairly  well.  Verrucaria  muraiis  is  considered  a  synonym  of  V.  rupestris 
Schrad.  in  Fink  (1935),  but  the  latter  is  considered  quite  different  by 
Zschacke,  with  an  endolithic,  sometimes  disappearing  thallus.  Zschacke 
states  that  V.  muraiis  is  found  on  sandstone  and  bricks. 

Distribution  — -  Arctic-boreal  element(?);  circumboreal. 

19.  Verrucaria  nigrescens  Pers.  Ann.  d.  Bot.  15:  36.  1795. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2827  (115). 

The  Long  Island  specimen,  together  with  one  collected  on  Nantucket 
Island  (Massachusetts)  (Brodo  3964  B),  agrees  in  most  respects  with 
descriptions  by  Zschacke  (1933)  and  Fink  (1935).  However,  the  black 
medullary  layer  mentioned  by  Zschacke  and  others  was  not  seen  in  all 
parts  of  the  thalli,  although  it  was  conspicuous  in  the  Nantucket  speci¬ 
men.  Servit  (1954)  describes  the  spores  of  this  species  as  20-28  x  1 1  p, 
but  these  measurements  disagree  with  those  of  all  previous  authors. 

Both  the  Long  Island  and  Nantucket  material  were  found  on  con¬ 
crete,  and  both  were  either  in  or  close  to  the  aerohaline  stratum  (p.  59). 
It  is  a  common  lichen  on  calcareous  rocks  in  Europe. 

Distribution  —  Connecticut,  Indiana,  Black  Hills,  Washington,  Mani¬ 
toba:  Temperate  element.  North  Temperate  subelement  (see  Fink,  1935); 
Europe;  Asia  ( Zahlbruckner,  1930). 

20.  Verrucaria  silicicola  Fink  in  Hedr.  Myco!ogia  25:  305.  1933. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2710  (111),  2826 

(115);  Three  Mile  Harbor,  Latham  32177,  April  16,  1951  (Latham); 
Orient,  Latham  36780,  April  14,  1950  (Latham);  Three  Mile  Harbor, 
East  Hampton,  Latham  36781.  April  19,  1949  (Latham);  Shelter  Island, 
Latham  36785,  June  1,  1944  (Latham);  Sag  Harbor,  Latham  36786, 
June  2,  1946  (Latham);  Orient,  Latham  (Holotype)  (MICH);  East 
Hampton,  Latham  2647,  April  20,  1926  (MICH);  East  Hampton,  Latham 
3995,  April  10,  1927  (MICH);  East  Hampton,  Latham  32177  (??  cf. 
above),  April  11,  1953  (MO). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


155 


This  species  is  similar  in  general  external  appearance  to  V.  micro- 
spora,  but  the  latter  has  much  smaller  perithecia  and  spores.  Both  species 
are  found  on  pebbles  and  small  stones  in  the  hydrohaline  stratum  in 
the  maritime  region  (figure  81). 

Distribution  —  Long  Island  (Fink,  1935):  Temperate  element, 
Maritime  subelement;  endemic. 

10.  DERMATOCARPON  Eschw. 

21.  Dermatocarpon  miniatum  (L. )  Mann,  Lich.  Bohm.  Obs.  Dispos. 
66.  1825.  Lichen  miniatus  L.  Sp.  PI.  1149.  1753. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk,  Latham  22242, 
May  6,  1926  (Latham).  The  specimen  was  found  on  a  rock  along 
railroad  tracks. 

Distribution  —  Massachusetts,  Connecticut,  Tennessee,  Alabama, 
Oklahoma,  Michigan,  Ontario,  Minnesota,  Black  Hills,  Washington, 
British  Columbia;  arctic  to  temperate  (Ahti,  1964):  Temperate  element, 
North  Temperate  subelement,  but  arctic  in  Asia  (Lynge,  1928),  Europe 
(  Lvnge,  1938),  and  Iceland  (Lynge,  1940a). 

PYRENULACEAE 

11.  PYRENULA  Ach. 

22.  Pyrenula  nitida  (Weig.)  Ach.  Syn.  Lich.  125.  1814.  Sphaeria 
nitida  Weig.  Obs.  Bot.  45.  1772. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25552  ( 52), 
Brodo  59-250  (61),  850  (41),  978  ( 63),  1221  (100a),  7657(88), 

1787  (127),  2210  (61),  2304  (93),  2539  (73),  2610  (84),  3232  (35), 
3320  (129);  Napeague,  Latham  2835,  March  1,  1927  (Latham);  Green- 
port,  Latham  3989,  April  1,  1927  (Latham). 

Pyrenula  nitida  is  the  indicator  species  of  a  well-known  and  well- 
studied  Fagus  community  in  Europe  (the  Pyrenuletum  nitidae  Hill.).  In 
both  Europe  and  Long  Island,  the  species  is  characteristic  of  smooth- 
barked  trees,  chiefly  Fagus  (and  Quercus  on  Long  Island),  in  moderately 
shaded  woods  (Barkman,  1958;  Almborn,  1948).  Its  position  in  a  Fagus 
community  continuum  seems  to  be  governed  by  light  availability  (Alm¬ 
born,  1948). 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Wisconsin,  Min¬ 
nesota;  throughout  the  United  States  (Fink,  1935);  Europe;  Asia  (Zahl- 
bruckner,  1930). 


12.  MELANOTHECA  Fee 

23.  Melanotheca  cruenta  (Mont.)  Mull.  Arg.  Bot.  Jahrb.  6:  397. 
1885.  Trypethelium  cruentum  Mont.  Ann.  Sci.  Nat.  II.  8:  537.  1837. 

Material  seen  —  SUFFOLK  COUNTY:  Gardiner’s  Island,  Latham, 
May  23,  1923  (Latham). 

A  description  and  discussion  of  this  species  can  be  found  in  lohn- 
son  (1959). 


156  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  Long  Island  specimen  extends  the  known  range  of  M.  cruenta 
slightly  northward.  This  range  extension  is  known  for  several  other 
coastal  plain  species,  among  them  Cladonia  santensis  and  Cladonia 
evansii.  The  specimen  was  found  on  a  tree  trunk  in  rich  woods. 

Distribution  —  Along  the  coastal  plain.  New  Jersey  to  Texas  (Fink, 
1935):  Temperate  element.  Coastal  Plain  subelement;  endemic. 

13.  TRYPETHELIUM  Spreng. 

24.  Trypethelium  virens  Tuck,  in  W.  Dari.  FI.  Cestr.  ed.  3,  453.  1853. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25735  (132), 

25743  (132),  25746  (132),  Brodo  59-194  (33),  2702  (111),  3070  (128), 
3201  (33),  3211  (33),  3254  (119);  Montauk,  Point  Woods,  Latham 
3992,  April  7,  1927  (Latham);  Orient,  Latham  Bros,  woods,  Latham 
3598C,  April  10.  1939  (Latham);  Napeague,  Latham  28356 A,  Febru¬ 
ary  9,  1 949  (  MO  ) . 

The  unusual  specificity  of  this  species  for  Ilex  spp.  and  Fagus  grandi- 
folia  was  discussed  on  p.  31.  One  specimen  (Latham  8598C )  was  col¬ 
lected  from  a  black  oak.  Johnson  (1959)  lists  a  number  of  other 
substrates  as  well.  Trypethelium  virens  is  apparently  skiophilous  (or 
hygrophilous?)  in  holly  groves,  thickets,  and  beech  forests;  it  is  never 
found  on  well-illuminated  trunks. 

Distribution  —  Trypethelium  virens  shows  an  unusual  North 
American  distribution  due  to  its  dual  substrate  specificity.  It  has  a 
typical  coastal  plain  distribution  from  Louisiana  through  Florida  to 
New  England  (Fink,  1935)  following  the  range  of  Ilex  opaca,  as  well 
as  an  Appalachian-Great  Lakes  distribution  following  the  range  of 
Fagus  grandifolia. 

Temperate  element,  Eastern  Temperate  subelement;  endemic. 

PORINACEAE 

14.  POR1NA  Mull.  Arg. 

25.  Porina  cestrensis  (Tuck,  in  W.  Dari.)  Mull.  Arg.  Flora  66:  338. 
1883.  Verrucaria  cestrensis  Tuck,  in  W.  Dari.  FI.  Cestr.  ed.  3.  452.  1853. 

Material  seen  - — -  SUFFOLK  COUNTY:  Orient  Point,  Latham  5, 
March  22,  1910  (NYS). 

The  type  material  of  this  species  ( Michener  204,  sub  Verrucaria 
cestrica )  was  examined  in  the  Farlow  herbarium.  A  diagnosis  of  the 
holotype  is  presented  in  tabular  form  in  the  discussion  of  P.  hibernica 
which  follows. 

Distribution  —  New  England  to  Georgia,  Alabama,  and  Tennessee 
(Fink,  1935):  Temperate  element.  Coastal  Plain  subelement;  endemic. 

26.  Porina  hibernica  P.  James  &  Swins.  in  Swins.  Lichenol.  2:  35. 
1962. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1783  B  (127), 
2598  (84),  3206  (33). 

This  species  bears  certain  resemblances  to  two  other  Porinae  from 
the  New  England  area,  P.  cestrensis  and  P.  rhaphidosperma  Mull.  Arg. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


157 


The  table  presented  below  points  out  some  of  the  differences  between 
them.  The  diagnoses  of  P.  centrensis  and  P.  rhaphidosperma  are  based  on 
the  type  specimens.  The  values  given  in  parentheses  under  P.  cestrensis 
are  measurements  of  other  specimens  which  were  studied. 


P.  cestrensis 

P.  rhaphidosprema 

P.  hibernica 

Thallus 

well  developed; 
greenish  black 

ashy  white  to  dirty 
green-grey;  smooth 
to  cracked,  well 
developed 

greenish  to 
olivaceous;  very 
thin,  smooth  or  scurfy 
to  almost  absent 

Perithecium 

0.15-0.25  mm 

0.20-0. 35mm 

0.20-0.35  mm 

Exciple 

hyaline 

carbonaceous 

carbonaceous 

Spores: 

size 

34-46  x  3-5  ;j. 
(30-42  x  5-6  u.) 

(63-)  100-120 
x  2-5  \j. 

58-65  x  5-7  ij. 

shape 

clavate,  straight 

acicular,  flexuous 

±  elongate-clavate 
to  ±  acicular, 
straight 

septa 

5-8 

(3-7  [-9]) 

(9-) 14-25 

(5-)  9-1 3  (-1 6) 

cell  size 

irregular 

equal 

frequently 

irregular 

Porina  hibernica  was  always  associated  with  P.  nuciila  on  oaks  in 
well  shaded  moist  woods. 

Distribution  —  First  North  American  record;  Ireland  (Kilarney: 
type  locality);  oceanic(?). 

27.  Porina  nucula  Ach.  Syn.  Meth.  Lich.  112.  1814. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1783  A  {111),  2598, 
sterile  (84),  3517  (33). 

This  species  is  one  of  the  few  Porinae  having  a  pale,  noncar- 
bonaceous  involucrellum. 

The  Long  Island  material  was  somewhat  aberrant  in  that  the  peri- 
thecia  were  small  and  did  not  have  the  “plaques”  or  lamellae  described 
by  Swinscow  (1962).  Herbarium  specimens  of  Porina  nucula  which 
I  examined  generally  had  a  smooth  to  verrucose  thallus,  but  the  Long 
Island  specimens  had  diffuse  coralloid  thalline  outgrowths.  These  out¬ 
growths  appear  like  Trentepohlia  filaments  which  have  partially  escaped 
lichenization.  Swinscow  (pers.  corr.)  said  that  these  specimens  were 
poorly  developed  but  otherwise  normal,  and  so,  perhaps  the  condition  is 
not  as  unusual  as  it  first  appears. 

Its  ecology  is  the  same  as  that  of  P.  hibernica. 

Distribution  —  Gulf  coastal  plain  (Fink,  1935):  Tropical  element 
(see  Swinscow,  1962),  Coastal  Plain  subelement;  Europe  (Swinscow, 
1962). 


158 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


GRAPHSDACEAE 

15.  XYLOGRAPHA  (Fr.)  Fr. 

28.  Xylographa  opegraphella  Will,  in  Rothr.  Proc.  U.S.  Nat.  Mus. 
7:  8.  1884. 

Material  seen  —  SUFFOLK  COUNTY:  Orient,  Latham  12,  May  1, 
1914  (Latham);  Orient,  Latham  1080,  April  1,  1915  (Latham). 

Norstictic  acid  was  demonstrated  in  KOH  from  the  specimen  with 
a  well-developed  thallus  (Latham  12)  and  this  substance  undoubtedly  is 
the  basis  of  the  KOH  +  red,  PD  +  yellow  reactions  in  the  medulla  of 
material  examined  by  Lamb  (1954).  The  other  Long  Island  specimen 
(Latham  1080)  had  almost  no  thallus  (but  was  identical  in  other  respects 
to  Latham  12)  and  norstictic  acid  could  not  be  demonstrated  in  the  very 
minute  thalline  particles  which  were  present. 

Xylographa  abietina  (Pers.)  Zahlbr.  differs  from  X.  opegraphella  in 
having  broader  spores  and  longer  lirellae.  An  exsiccat  collection  of 
X.  abietina  (California  Fungi  no.  850)  had  spores  12-14  x  6-7  rj.  with 
lirellae  up  to  a  millimeter  or  more  long.  No  norstictic  acid  could  be  found 
in  the  specimen,  which  had  virtually  no  thallus. 

Xylographa  opegraphella  is  confined  to  old  wood. 

Distribution  —  New  England  coast  (Fink,  1935),  Nova  Scotia; 
Alaska  (Rothrock,  1884;  Cummings,  1910):  Temperate  element,  Oceanic 
subelement;  endemic. 


16.  GRAPHiS  Adans. 

29.  Graphis  scripta  (L.)  Ach.  Kgl.  Vet.  Akad.  Nya  Handl.  145. 
1809.  Lichen  scriptus  L.  Sp.  PI.  1140.  1753. 

Material  seen  —  SUFFOLK  COUNTY:  52  specimens  collected  by 
Imshaug  and/or  Brodo;  12  specimens  collected  by  Latham  (Latham). 

The  lirellae  of  this  species  are  extremely  variable  in  length,  breadth, 
and  degree  of  branching.  Gross  lirelline  characters  are  therefore  of  little 
use  in  defining  the  species. 

Graphis  scripta  is  common  on  the  bark  of  various  deciduous  trees, 
usually  in  partial  shade,  and  is  mainly  associated  with  the  red  oak  forest 
on  the  north  shore  (figure  61). 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Massachusetts, 
North  Carolina,  Tennessee,  Michigan,  Wisconsin,  Indiana,  Minnesota, 
Washington,  Alaska:  Temperate  element.  North  Temperate  subele- 
ment(?);  Europe:  Asia  (Vainio,  1928;  Zahlbruckner,  1930b). 

17.  PHAEOGRAPHIS  Mull.  Arg. 

30.  Phaeographis  dendritiea  (Ach.)  Mull.  Arg.  Flora  65 :  382.  1882. 
Opegrapha  dendritiea  Ach.  Meth.  Lich.  31.  pi.  1,  f.  10.  1803. 

Material  seen  —  NASSAU  COUNTY:  Brodo  1509  ( 14),  546  ( 12), 
554  (12).  SUFFOLK  COUNTY:  72  specimens  collected  by  Imshaug 
and/or  Brodo;  12  specimens  collected  by  Latham  (Latham);  Greenport, 
Latham  31919,  April  12,  1953  (MO). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  159 

The  species  is  found  on  the  bark  of  various  deciduous  trees  in  well- 
lighted  or  partially  shaded  woods. 

Distribution  —  Eastern  United  States  (Fink,  1935);  Temperate 
element,  East  Temperate  subelement;  Europe;  Asia  (Zahlbruckner, 
1930b). 


DIPLOSCHISTACEAE 

18.  DIPLOSCHISTES  Norm. 

31.  Diploschistes  scruposus  (Schreb.)  Norm.  Nyt.  Mag.  Naturv.  7: 
232.  1853.  Lichen  scruposus  Schreb.  Spic.  FI.  Lips.  133.  1771. 
var.  scruposus 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3847  (76);  Orient, 
Latham,  October  4,  1917  (Latham);  Sag  Harbor,  Latham,  May  10 
1924  (Latham). 

var.  parasiticus  (Sommerf.)  Zahlbr.  Cat.  Lich.  Univ.  2:  672.  1924. 

Lecanora  scruposa  var.  parasitica  Sommerf.  Suppl.  FI.  Lapp.  100. 
1826. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-172  (100B). 

All  the  specimens  were  on  noncalcareous  rock,  except  var.  parasiticus 
which  was  collected  on  the  sterile  squamules  of  a  species  of  Cladonia. 
Diploschistes  scruposus  was  treated  by  Fink  (1935)  in  the  genus  Urceo- 
laria. 

Distribution  —  Maine,  Connecticut,  Michigan,  Oklahoma,  Arizona, 
Black  Hills,  Washington,  Manitoba,  Baffin  Island:  Arctic-boreal  element; 
circumboreal. 


GYALECTACEAE 

19.  DIMERELLA  Trev. 

32.  Dimerella  diluta  (Pers.)  Trev.  Rend.  Reale  1st.  Lomb.  Sci.  13: 

65.  1880.  Peziza  diluta  Pers.  Syn.  Meth.  Fung.  668.  1801. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3200  (33). 

Both  this  species  and  the  one  following  were  treated  under  the  genus 
Microphiale  by  Fink  (1935). 

The  Long  Island  specimen  was  found  on  the  bark  of  an  old  oak  in 
the  dense  shade  of  an  Ilex  opaca  grove  on  Fire  Island. 

Distribution  — -  Eastern  United  States  (Fink,  1935);  Maine,  North 
Caro  na,  B’ack  Hills,  Saskatchewan:  Temperate  element.  East  Temperate 
sube!ement(?) ;  Europe;  Asia  (Vainio,  1928). 

33.  Dimerella  lutea  (Dicks.)  Trev.  Rend.  Reale  1st.  Lomb.  Sci.  13: 

66.  1880.  Lichen  luteus  Dicks,  Fasc.  PI.  Crypt.  Brit.  1:  11,  pi.  2,  f.  6. 
1785. 

Material  seen  —  SUFFOLK  COUNTY:  Orient,  Latham  1087,  May 
3,  1914  (Latham). 

A  specimen  of  this  species  was  also  found  on  Cape  Cod,  (Massachu¬ 
setts)  in  a  bog  ( Brodo  4323B). 


160  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Distribution  —  Eastern  United  States,  and  Canada  (Fink,  1910); 
Maine,  North  Carolina,  Washington:  Temperate  element,  East  Tem¬ 
perate  subelement(  ?) ;  Europe;  Asia  (Vainio,  1928). 

COLLEMATACEAE 

20.  COLLEMA  Wigg. 

34.  Collema  subfurvum  (Mull.  Arg. )  Degel.  Bot.  Not.  139.  1948. 
Synechoblastus  flaccidus  v.  subfurvus  Miill.  Arg.  Proc.  Roy.  Soc.  Edinb. 
11:  457.  1882. 

Material  seen  —  QUEENS  COUNTY:  Jamaica,  G.  B.  Brainerd, 
1866  (BKL);  Jamaica,  G.  B.  Brainerd,  1866,  ( BKL  031870).  SUFFOLK 
COUNTY:  Orient,  Latham  787 A,  May  3,  1914  (Latham);  Napeague, 
Latham  2845,  March  1,  1927  (Latham);  Montauk,  Latham  28309,  Febru¬ 
ary  9,  (Latham);  Shelter  Island,  Latham  36949,  May  4,  1943  (Latham). 

Collema  subfurvum  differs  from  closely  related  C.  flaccidum  (Ach.) 
Ach.  (Syn.  Synechoblastus  rupestris  Trev.)  in  having  globular  rather  than 
squamiform  isidia,  and  in  its  corticolous  rather  than  saxicolous  substrate 
preference  (Degelius,  1954).  It  is  usually  found  on  oak  bark. 

Distribution  —  New  England,  Smoky  Mountains  (Tennessee),  Iowa, 
Illinois  (Degelius,  1954):  Temperate  element,  Appalachian  subelement, 
Appalachian-Great  Lakes  unit;  Europe  (oceanic  localities)  and  Asia 
(Degelius,  1954). 

21.  LEPTOGIUM  S.  Gray 

35.  Leptogium  corticola  (Tayl.)  Tuck,  in  Lea,  Cat.  PI.  Cine.  47. 
1849.  Collema  corticola  Tayl.  J.  Bot.  5:  195.  1847. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk,  Latham  3993 
(p.p.),  April  6,  1927  (Latham). 

Degelius  (1940)  discusses  the  nomenclatural  problems  pertaining  to 
this  species.  The  Long  Island  material  agrees  with  the  original  descrip¬ 
tion  as  well  as  Degelius’  additions  to  it.  Sierk  (1964)  presents  a  detailed 
discussion  of  the  species. 

Distribution  —  Temperate  element,  East  Temperate  subelement, 
Adriatic  coast  in  Europe  (map:  Sierk,  1964). 

36.  Leptogium  cyanescens  (Ach.)  Korb.  Syst.  Lich.  Germ.  420. 
1855.  Collema  tremelloides  v.  cyanescens  Ach.  Syn.  Meth.  Lich.  326. 
1814.  non  Lichen  cyanescens  Pers.  or  Parmelia  cyanescens  Ach.  (De¬ 
gelius,  1935). 

Material  seen  — -  QUEENS  COUNTY:  Jamaica,  G.  B.  Brainerd, 
1866?  (BKL).  SUFFOLK  COUNTY:  Brodo  2126  (102);  Orient,  Latham 
787,  May  3,  1914,  (Latham);  Orient,  Latham  8199,  April  16,  1928 
(Latham);  Greenport,  Latham  8618,  June  1,  1931  (Latham);  North  Sea, 
Latham  23333,  March  26,  1954  (Latham);  Montauk,  Latham  28309 A, 
February  9,  1949  (Latham);  Three  Mile  Harbor,  Ogden  5406,  May  11, 
1954  (NYS). 

This  species  was  apparently  included  in  L.  tremelloides  (L.)  S.  F. 
Gray  by  Fink  (1935).  Leptogium  tremelloides,  however,  is  strictly  an 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


161 


Old  World  species  (Sierk,  1964).  The  confusing  nomenclature  of 
L.  cyanescens  has  been  clarified  by  Degelius  (1935).  The  species’  oceanic 
affinities  are  noted  by  Degelius  (  1935  and  1941).  The  distribution  of 
L.  cyanescens  on  Long  Island  (figure  31),  showing  a  restriction  to  the 
foggy  eastern  tip,  reflects  these  oceanic  requirements.  Sierk  (1964)  dis¬ 
cusses  its  morphology,  ecology,  and  distribution  in  detail. 

On  Long  Island,  the  species  is  usually  found  on  mossy  tree  bases. 

Distribution  —  Temperate  element,  basically  East  Temperate  sub¬ 
element,  with  scattered  occurrences  in  the  Black  Hills,  western  Canada 
and  coastal  Alaska,  Europe,  Asia  (map:  Sierk,  1964). 

PANNARIACEAE 

22.  PLACYNTHIUM  S.  Gray 

In  her  recent  North  American  monograph  of  the  genus,  Henssen 
(1963)  placed  Placynthium  into  the  Peltigeraceae  based  on  ascocarp 
development.  Since  Henssen’s  revision  of  the  cyanophycean  lichens  and 
their  families  is  still  not  complete,  the  older  family  concepts  will  be 
retained  for  the  time  being. 

37.  Placynthium  nigrum  (Huds.)  S.  Gray  Nat.  Arr.  Brit.  PI.  395. 
1821.  Lichen  niger  Huds.  FI.  Angl.  ed.  2,  2:  524.  1778. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3921  (54). 

Although  P.  nigrum  is  considered  squamulose  or  even  subfoliose  by 
some  authors,  the  Long  Island  material  was  all  crustose,  occasionally  form¬ 
ing  small  subsquamulose  areoles.  Henssen  (1963)  presents  a  detailed 
account  of  the  species’  morphology  and  development. 

This  inconspicuous  species  is  probably  more  abundant  than  the  col¬ 
lection  records  show.  It  was  found  in  a  shaded  woods  on  old  concrete 
foundations. 

Distribution  —  Arctic-boreal  element  (map:  Henssen,  1963); 
Europe;  Asia  (Lynge,  1928). 

23.  PANNARIA  Del. 

38.  Pannaria  lurida  (Mont.)  Nyl.  Mem.  Soc.  Sci.  Nat.  Cherb.  5: 
109.  1857.  Collema  luridum  Mont.  Ann.  Sci.  Nat.  II.  18:  236.  1842. 

Material  seen  —  SUFFOLK  COUNTY :  Montauk  Woods  north  of 
Fresh  Pond,  Latham  28322,  February  9,  1949  (Latham);  Orient  Point, 
Latham  5,  April  4,  1910  (NYS).  COUNTY  UNKNOWN:  Long  Island 
(?),  Austin  (BKL  031953). 

The  species  was  found  on  oak  and  red  cedar  bark. 

Distribution  —  Eastern  United  States  (Fink,  1935):  Tropical  Ele¬ 
ment  (Zahlbruckner,  1925),  Appalachian-Temperate  subelement. 

STICTACEAE 

LOBARIA  Schreb. 

39.  Lobaria  pulmonaria  (L.)  Hoffm.  Deutschl.  FI.  2:  146.  1796. 
Lichen  puhnonarius  L.  Sp.  PI.  1145.  1753. 


162  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Material  seen  —  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 
1867  (BKL  031881).  SUFFOLK  COUNTY:  Brodo  887  (56),  1021 
(112),  1045  (112),  2154  (102);  12  specimens  collected  by  Latham 
(Latham) . 

Fink  (1935)  treated  L.  puhnonaria  under  the  genus  Sticta. 

This  species  shows  some  variation  in  isidia  and  soredia  production 
in  various  parts  of  its  range.  The  Long  Island  specimens  all  have 
isidiate-soralia  on  the  thallus  margins  and  ridges,  hut  they  may  be  more 
common  in  some  individuals  than  in  others.  The  granular  soredia  are 
sometimes  hard  to  see  until  most  of  the  isidia  have  fallen  away.  The 
isidia  vary  from  being  short,  almost  like  papillae,  to  elongate  cylindrical, 
and  finally  coralloid. 

The  species  is  rare  on  Long  Island.  It  is  confined  to  tree  bases  in 
the  oceanic  areas  of  the  eastern  tip  of  the  island  and  bog  trees  (especi¬ 
ally  Acer  rubrum)  outside  this  area. 

Degelius  (1935,  p.  223)  stated  that  L.  puhnonaria  favors  an  oceanic 
climate  but  is  not  restricted  to  an  oceanic  distribution. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Tennessee,  Michi¬ 
gan,  Ontario,  Indiana,  Washington,  British  Columbia,  Alaska:  Tem¬ 
perate  element.  North  Temperate  subelement  (Appalachian-Great  Lakes: 
Hale,  1961a);  Europe;  Asia  (Zahlbruckner,  1930;  Magnusson,  1940). 

40.  Lobaria  quercizans  Michx.  FI.  Bor-Amer.  2:  324.  1803. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  (Brainerd  ?) ,  1867 

(BKL  031874).  SUFFOLK  COUNTY:  Brodo  1040  ( 112),  2145  (102), 
2801  (102);  Napeague,  Latham  2858,  March  1,  1927  (Latham); 
Napeague,  Latham  2837,  March  1,  1927  (Latham);  Montauk,  Latham 
28307,  February  9,  1949  (1945?)  (Latham,  MO);  Riverhead,  Latham 
36869,  May  16,  1960  (Latham);  Montauk,  Latham  36884,  April  4,  1949 
(Latham);  Jamesport,  Latham  36948,  April  19,  1951  (Latham);  (no 
locality),  Latham,  May  6,  1920  (Latham);  Eastport,  Schrenk,  June  28, 
1894  (MO). 

Lobaria  quercizans  is  the  North  American  vicariad  of  L.  amplissima 
(Scop.)  Forss.  (Degelius,  1940;  Hale,  1957a),  a  well  known  European 
oceanic  species  (see  Degelius,  1 935 ) .  Lobaria  quercizans  was  considered 
under  the  latter  name  in  Fink  (1935).  The  North  American  species  also 
appears  to  have  an  oceanic  distribution  (Degelius  1941),  and  on  Long 
Island  is  restricted  to  the  fog  belt  and  bogs.  It  often  is  found  associated 
with  Lobaria  puhnonaria. 

Distribution  —  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit  (map:  Hale,  1957a);  endemic. 

NEPHROMACEAE 

25.  NEPHROMA  Ach. 

41.  Nephroma  laevigatum  Ach.  Svn.  Lich.  242.  1814.  non  auct. 

Material  seen  —  SUFFOLK  COUNTY :  Montauk,  Latham  36784, 

May  6,  1929  (Latham). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


163 


A  full  discussion  of  the  taxonomy  and  distribution  of  this  species 
was  presented  by  Wetmore  (1960).  Long  Island  is  the  southernmost 
locality  for  the  species  on  the  east  coast.  The  specimen  was  found  on  rock. 

Distribution  —  East  and  west  coasts  of  North  America:  Temperate 
element,  Oceanic  subelement  (Wetmore,  1960);  oceanic  regions  of  Europe 
( Degelius,  1935);  Asia  (Vainio,  1928). 

PELTIGERACEAE 

26.  SOLORINA  Ach. 

42.  Solorina  saccata  (L.)  Ach.  Kgl.  Vet.  Akad.  Nya  Handl.  228. 
1808.  Lichen  saccatus  L.  FI.  Suec.  ed.  2,  419.  1755. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk,  Latham  36883, 
October  7,  1926  (Latham). 

The  specimen  was  collected  on  a  rocky  bank. 

Distribution  —  Michigan,  Ontario,  Minnesota,  Black  Hills,  Wash¬ 
ington,  Alaska,  Manitoba,  Quebec,  Baffin  Island:  Arctic-boreal  element; 
circumboreal. 


27.  PELTIGERA  Willd. 

43.  Peltigera  aphthosa  (L.)  Willd.  FI.  Berol.  Prodr.  347.  1787. 
Lichen  aphtosus  L.  Sp.  PI.  1148.  1753. 

var.  variolosa  (Mass.)  Thoms.  Trans.  Wise.  Acad.  Sci.  38:  253.  1947. 
Peltigera  aphthosa  f.  variolosa  Mass.  Sched.  Crit.  Ill:  64.  1856. 
Material  seen  —  KINGS  COUNTY :  New  Lots,  G.  B.  Brainerd 
( BKL  031888).  SUFFOLK  COUNTY:  Gardiner's  Island,  Latham 
September  22,  1922  (Latham);  Fisher’s  Island,  Latham ,  June  24,  1929 
(Latham);  Montauk,  Latham,  May  17,  1942  (Latham). 

The  dark  veins  on  the  lower  surface  of  this  variety  distinguish  it 
from  var.  aphthosa  (var.  typica  in  Thomson,  1950a). 

The  material  is  from  the  ground  in  dry  woods,  and  from  a  rock 
(figure  70). 

Distribution  —  Arctic-boreal  element,  circumboreal  (map:  Thomson, 
1950a). 

44.  Peltigera  canina  (L.)  Willd.  FI.  Berol.  Prodr.  347.  1787.  Lichen 
caninus  L.  Sp.  PI.  1149.  1753. 

var.  rufescens  (Weiss)  Mudd,  Man.  Brit.  Lich.  82.  1861. 

Lichen  caninus  var.  rufescens  Weiss,  PI.  Crypt.  FI.  Goet.  79.  1770. 
Material  seen  —  NASSAU  COUNTY:  Massapequa,  S.  Cain  188, 
July  7,  1935  (NY).  SUFFOLK  COUNTY:  Devon,  Latham,  May  2,  1955 
(Latham);  Three  Mile  Harbor,  Latham  27207,  April  17,  1947  (Latham); 
Napeague,  north  of  Fresh  Pond,  Latham  8118,  April  6,  1938  (Latham); 
Three  Mile  Harbor,  Hands  Creek,  Latham  2646,  April  20,  1926  (La¬ 
tham);  Napeague,  Latham  36978,  May  3,  1947  (Latham). 

Following  Thomson  (1950a),  three  varieties  of  this  species  can  he 
recognized  as  occurring  on  Long  Island. 


164  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  questionable  taxonomic  rank  of  var.  rufescens  is  discussed  on 
p.  110-111.  On  Long  Island  it  is  relatively  rare,  occurring  mainly  in 
dry  woods  on  tree  bases. 

var.  spuria  (Ach.)  Schaer.  Lich.  Helvet.  Spicil.  6:  265.  1833. 

Lichen  spurius  Ach.  Lich.  Suec.  Prodr.  159.  1798. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-280  (53),  2291 
(87);  Northwest,  Latham  26133,  April  10,  1947  (Latham). 

Variety  spuria  has  only  been  collected  in  its  sorediate  stage  on 
Long  Island.  It  has  been  clearly  established  that  the  sorediate  form  is  a 
juvenile  stage  of  var.  spuria  (Dahl,  1950).  Latham  26133  has  both  apo- 
thecia  and  soredia  and  appears  similar  to  var.  rufescens,  which  in  turn 
is  said  to  intergrade  with  var.  canina  (var.  albescens)  (Thomson,  1950a). 

It  was  found  on  dry,  sandy  soil. 

var.  ulorrhiza  (Florke)  Schaer.  Enum.  Crit.  Lich.  Europ.  20.  1850. 

Pelticlea  ulorrhiza  Florke,  Deutsch.  Lich.  no.  154.  1821. 

Material  seen  —  SUFFOLK  COUNTY:  Riverhead,  Latham,  No¬ 
vember  1,  1913  (Latham). 

The  Latham  specimen  was  found  on  the  ground  in  a  dry  woods. 

Distribution  (of  all  varieties)  —  Arctic-boreal  element;  circum- 
boreal  (maps:  Thomson,  1950a). 

45.  Peltigera  polydactyla  (Neck.)  Hoffm.  Desc.  Adumbr.  PI.  Lich. 
1:  19,  pi.  4,  f.  1.  1790.  Lichen  polydactylon  Neck.  Meth.  Muse.  85.  1771. 

Material  seen  —  QUEENS  COUNTY:  Jamaica,  G.  B.  Brainerd, 
May  1866  ( BKL  031889).  SUFFOLK  COUNTY:  Montauk,  Latham, 
May  12,  1920  (Latham). 

Peltigera  polydactyla  is  most  closely  related  to  P.  horizontalis 
(Huds.)  Baumg.,  from  which  it  is  distinguished  by  its  vertically-oriented 
apothecia  and  its  longer  and  narrower  spores.  Thomson  (1950a)  reports 
the  spores  of  the  latter  species  to  be  24-45  x  3.5-6  [j.. 

The  Long  Island  material,  having  broad,  conspicuous  veins  on  the 
lower  thallus  surface,  represents  var.  polydactyla  (var.  typica  of  Thom¬ 
son,  1950a). 

Latham's  specimen  was  found  at  the  base  of  a  tree  in  an  oak  woods. 

Distribution  —  Arctic-boreal  element  (map:  Thomson,  1950a),  also 
Baffin  Island,  Manitoba;  circumboreal. 

46.  Peltigera  praetextata  (Florke  in  Somm.)  Vain.  Termeszetr. 
Fuzetek  22:  306.  1899.  Peltidea  ulorrhiza  var.  praetextata  Florke  in 
Somm.  Suppl.  FI.  Lappon.  123.  1826. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  7049(112),  1254 
(48),  2041  (45),  2133  (102),  2469  (23);  Greenport,  Latham  53,  May 
10,  1914  (Latham);  Three  Mile  Harbor,  Latham,  November  21,  1926 
(Latham);  Riverhead,  Latham,  February  2,  1923  (Latham);  16  speci¬ 
mens  collected  by  Latham  (Latham). 

This  species  is  very  similar  to  P.  canina  var.  rufescens  and  seems 
to  differ  only  in  its  ability  to  produce  regeneration  squamules  on  the 
thallus  surface  and  margins.  Experiments  on  the  production  of  isidia 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  165 

(regeneration  squamules)  were  performed  by  Thomson  (1948)  and 
Lindahl  (1953)  and  resulted  in  two  entirely  opposite  points  of  view 
regarding  the  taxonomic  value  of  the  structures.  Thomson  found  that 
wounded  thalli  of  P.  canina  var.  rufescens  regenerate  on  some  lobes  and 
not  on  others,  whereas  Lindahl  found  that  no  thalli  of  P.  canina  sens.  str. 
regenerated  and  only  P.  praetextata  showed  regeneration.  It  is  possible  that 
Thomson  was  working  with  true  P.  praetextata  “hidden”  by  its  original 
lack  of  regeneration  squamules  (which  Lindahl  concedes  may  happen) 
and  that  true  P.  canina  sens.  str.  would  not  produce  regeneration  even 
in  the  United  States.  It  is  also  possible  that  only  under  certain  condi¬ 
tions  will  species  other  than  praetextata  regenerate  and  these  conditions 
were  met  in  Wisconsin  and  not  in  Sweden,  or  that  the  America  popula¬ 
tions  of  P.  canina  differ  in  regeneration  properties  from  the  European 
populations,  a  difference  which  might  be  of  taxonomic  importance. 

This  entire  problem,  as  it  appears  to  me,  is  far  from  settled  and 
should  be  investigated  further.  Until  more  work  is  done,  however,  the 
European  concept  of  P.  praetextata  will  be  accepted. 

The  species  is  most  frequently  found  growing  on  mossy  tree  bases 
in  oak  woods. 

Distribution  —  Arctic-boreal  element  (map:  Thomson,  1950a); 
Europe;  Asia  (Magnusson,  1940). 

LECIDEACEAE 

28.  LECIDEA  Ach. 

47.  Lecidea  aeruginosa  Borr.  in  Hook,  and  Sowerb.  Suppl.  Engl. 
Bot.  1:  tab.  2682.  1831.  Lecidea  flexuosa  (Fr. )  Nyl.  Act.  Soc.  Linn.  Bord. 
21:  356.  1856. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25834  (86), 
Brodo  657  (79),  1157  (70),  1612  (69),  2213  (61),  2333  (44),  2548 
(73),  2732  (111),  2964  (95),  3319  (129),  3336  (18),  2536  (49);  Orient, 
Latham  38174,  December  15,  1964  (Latham). 

The  separation  of  sterile  material  of  L.  aeruginosa  from  L.  botryosa 
is  discussed  under  the  latter  species.  Laundon  (1962)  regarded  L.  aeru¬ 
ginosa  (sub  L.  flexuosa)  as  synonomous  with  L.  granulosa.  On  Long 
Island,  however,  except  for  spore  size,  the  two  are  not  at  all  similar  either 
morphologically  or  ecologically.  Lecidea  aeruginosa  has  black  or  lead- 
colored  plane  apothecia,  each  with  a  thin  hyaline  hypothecium;  L.  granu¬ 
losa  has  large,  brown,  irregularly  convex  to  almost  hemispherical  apothecia, 
each  with  a  thick  opaque  hypothecium.  In  addition,  the  former  species  is 
restricted  to  lignum  and  the  latter  is  found  only  on  sandy  soil.  I  have 
examined  material  from  the  Black  Hills  of  South  Dakota,  where  both 
species  occur  on  old  wood,  and  still  the  fertile  material  of  L.  aeruginosa 
is  easily  distinguished  from  L.  granulosa. 

Distribution  —  Connecticut,  Minnesota,  Black  Hills;  throughout  the 
United  States  (Fink,  1935):  Temperate  element.  North  Temperate  sub¬ 
element  (?);  Europe;  Asia  (Vainio.  1928). 


166  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

48.  Lecidea  albocaerulescens  (Wulf.  in  Jacq.)  Ach.  Meth.  Lich.  52. 
1803.  Lichen  albocaerulescens  Wulf.  in  Jacq.  Collect.  Bot.  2:  184,  f.  1. 
1788. 

Material  seen  —  NASSAU  COUNTY:  Brodo  549  (12).  SUFFOLK 
COUNTY:  Brodo  2166  (99),  2434  (20),  2580  (96),  2689  (110),  2743 
(111),  3019  (17),  3034  (50),  3037  (50),  3119  (34),  3272  (119), 
3274  (119),  3411  (134),  3875  (62);  Gardiner’s  Island,  Latham,  March 
30,  1921  (Latham);  Greenport,  Latham  3966,  April  1,  1927  (Latham); 
Greenport,  Latham  39,  May  10,  1914  (Latham);  Three  Mile  Harbor. 
Latham  32651,  May  25,  1954  (Latham). 

This  striking  saxicolous  species  is  easily  identified  in  the  field  by 
its  pruinose  apothecia,  dark  apothecial  margins,  and  smooth  grey  thallus. 
All  but  one  specimen  on  Long  Island  were  shown  to  contain  stictic  acid 
(both  by  paper  chromatography  and  recrystallization  in  GAoT  solu¬ 
tion).  The  exception  ( Brodo  549)  contained  norstictic  acid  (red  acicular 
crystals  in  KOH).  This  stictic-norstictic  shift  is  a  common  phenomenon 
in  lichen  chemistry  and  can  he  seen  in  Lecanora  cinerea  and  several 
species  of  Parmelia.  Norstictic  acid  has  never  been  reported  for  this 
species  before. 

Lecidea  albocaerulescens  is  narrowly  restricted  to  shaded  granitic 
rocks  and  is  only  found  in  the  poorly  lighted,  red  oak  forests  of  the 
north  shore  (figure  59). 

Distribution  —  Connecticut,  Tennessee,  Michigan,  Indiana,  Okla¬ 
homa,  Minnesota,  Washington,  Alaska;  Eastern  United  States  and  Wash¬ 
ington  (Fink,  1935);  Temperate  element.  North  Temperate  subelement; 
Europe:  Asia  (Lynge,  1928). 

49.  Lecidea  anthracophila  Nyl.  Flora  48:  603.  1865. 

Material  seen  —  SUFFOLK  COUNTY:  23  specimens  collected 
by  Imshaug  and/or  Brodo. 

Fink  (1935)  lists  this  species  with  the  genus  Psora. 

The  PD  +  red  constituent  of  L.  anthracophila  is  apparently  funtar- 
protocetraric  acid  hut  does  not  show  exactly  the  same  Rr  value  as  known 
fumarprotocetraric  acid  (as  in  Cladonia  subtenuis)  in  paper  chroma¬ 
tography  (solvent:  pyridine,  ethyl  acetate,  and  water).  Fumarprotocetraric 
acid  usually  has  an  Rf  of  approximately  0.30  to  0.45  and  the  Lecidea 
anthracophila  material  has  an  of  approximately  0.40  to  0.55.  In  all 
other  characters  (co'or  reaction  with  PD,  fluorescence  in  UV  before  and 
after  reaction  with  PD,  etc.)  it  is  identical  to  fumarprotocetraric  acid. 

The  species  is  found  only  on  fresh  or  charred  bark  of  Pinus  rigida 
(see  page  49);  figure  42).  Fink  (1935)  reported  it  from  old  wood. 

Distribution  —  Vermont,  Massachusetts,  New  Jersey,  and  North 
Carolina  (Fink,  1935):  Temperate  element.  East  Temperate  subelement; 
Europe. 

50.  Lecidea  botryosa  (Fr.)  Th.  Fr.  Lich.  Scand.  1:  454.  1874. 
Biatora  botryosa  Fr.  Kgl.  Vet.  Akad.  Nya  Handl.  268.  1822. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


167 


Material  seen  —  NASSAU  COUNTY:  Brodo  3494  (4).  SUFFOLK 
COUNTY:  Imshaug  25633  (NW  of  29),  25636a  (NW  of  29),  Brodo 
3202  (33). 

When  steri'e,  this  species  closely  resembles  L.  aeruginosa  which, 
however,  is  C  +  red.  If  apothecia  are  present,  the  hypothecial  color 
(hyaline  in  L.  aeruginosa  and  brown  in  L.  botryosa)  distinguishes  the 
two. 

The  species  is  almost  entirely  restricted  to  old  wood.  It  was  found 
once  (Brodo  3494)  growing  on  the  base  of  an  old  black  oak  in  a  shaded 
woods. 

Distribution  —  Michigan,  Arizona,  Manitoba;  Adirondack  Moun¬ 
tains  of  New  York,  New  Hampshire,  with  doubtful  occurrences  in  the 
west  coast  (Lowe,  1939):  Temperate  element.  North  Temperate  sub- 
e'ement;  northern  Europe,  Asia  (Lowe,  1939). 

51.  Leeidea  coarctata  (Turn,  in  Srn.  and  Sowerby)  Nyl.  Act.  Soc. 
Linn.  Bord.  21:  358.  1856.  Lichen  coarctatus  Turn,  in  Sm.  and  Sowerby 
Engl.  Bot.  8:  pi.  534.  1799. 

Material  seen  — •  QUEENS  COUNTY:  Brodo  525  (3).  SUFFOLK 
COUNTY:  Brodo  59-308  (54),  59-310  (54),  791  (90A),  2342  (44), 
2688  (110),  1782  (127),  2531  (49),  2720  (111),  3901  (112); 

Orient,  Latham ,  March  18,  1914  (Latham);  Shelter  Island,  Latham 
22177 ,  October  26,  1944  (Latham);  Montauk,  Latham  28127,  October  8. 
1954  (Latham). 

This  is  the  only  species,  found  on  pebbles,  which  has  small  brown 
apothecia.  The  white,  areolate,  C  +  red  thallus  add  to  its  distinctiveness. 
Leeidea  coarctata  is  often  associated  with  L.  erratic  a  and  Rhizocarpon 
obscuratum  on  pebbles  and  small  stones. 

Distribution  — •  Nova  Scotia,  Maine,  Connecticut,  Indiana,  Min¬ 
nesota,  British  Columbia;  northern  United  States  (Fink,  1935):  Tem¬ 
perate  element,  North  Temperate  subelement;  Europe. 

52.  Leeidea  eyrtidia  Tuck.  Proc.  Amer.  Acad.  Arts  Sci.  12:  181. 
1877. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1684  (88),  2330 
(44),  2697  (110),  3078A  (128),  3120  (34),  3125  (34),  3287  (119), 
3903  (112);  Greenport,  Latham  3974,  April  1,  1927  (Latham);  Shelter 
Island,  Latham  22177,  October  26,  1944  (Latham);  Shelter  Island, 
LaOiam  2 2879 A,  October  26,  1944  (Latham);  She'ter  Island,  Latham 
22880,  October  26,  1944  (Latham);  Shelter  Island,  Latham  22880, 
October  26,  1944  (Latham);  Latham  31015,  February  2,  1940  (Latham), 
Mon*auk  Point,  Latham,  April  12,  1956  (Latham). 

Leeidea  eyrtidia  is  superficially  very  similar  to  L.  erratica.  However, 
the  epithecium  and  the  outer  portions  of  the  exciple  are  greenish  black 
in  the  latter  and  brown  in  the  former  species.  Magnusson  (1952)  de¬ 
scribed  L.  nearingii  which,  from  its  description,  appears  very  similar 
to  L.  eyrtidia.  Leeidea  nearingii  has  a  brown-black  thallus,  whereas  L. 
eyrtidia  has  a  pale  to  dark  brownish-green  thallus.  The  thalli  of  both 


168  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

species  are  thin  and  continuous.  The  distinctions  are  therefore  very 
questionable  from  the  published  descriptions,  but  since  the  type  of 
L.  nearingii  has  not  been  examined  no  further  conclusions  can  be  made 
concerning  its  validity  as  a  species. 

One  specimen  had  much  larger  apothecia  than  any  of  the  others, 
but  agreed  in  other  respects  with  the  descriptions  of  L.  cyrtidia. 

The  species  is  common  on  pebbles  and  small  stones  in  dry  woods 
or  fields. 

Distribution  —  Eastern  United  States  (Lowe,  1939):  Temperate 
element,  East  Temperate  subelement;  endemic. 

53.  Lecidea  erratiea  Kdrb.  Parerg.  Lich.  223.  1861. 
var.  erratiea 

Materia!  seen  —  QUEENS  COUNTY:  Brodo  524  (3).  NASSAU 
COUNTY:  Brodo  545  (12),  3506  (10).  SUFFOLK  COUNTY:  32 
specimens  collected  by  Imshaug  and/or  Brodo;  Shelter  Island,  Latham 
22879B,  October  26,  1944  (Latham);  Shelter  Island,  Latham  22883, 
October  26,  1944  (Latham);  Riverhead,  Latham  24271,  March  16, 
1946  (Latham);  Quogue,  Latham  28254  (Latham);  Montauk  Point, 
Latham  29305,  May  6,  1949  (Latham);  Orient,  Brown  Brothers  Site, 
Latham  29928,  November  4,  1951  (Latham);  Riverhead,  North  River, 
Latham  34271,  March  16,  1946  (Latham);  East  of  Sag  Harbor,  Latham, 
October  19,  1945  (Latham). 

var.  planetica  (Tuck.)  Lowe,  Lloydia  2:  279.  1939. 

Lecidea  planetica.  Tuck.  Syn.  N.  Am.  Lich.  2:  131.  1888. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3012  (17). 

Magnusson  (1936)  recognized  several  species  as  being  closely 
related  to  L.  erratiea :  L.  sylvicola  Flot.,  L.  cyrtidia  Tuck.,  L.  micytho 
Tuck.,  and  L.  planetica  Tuck.  Lowe  ( 1939)  whose  work  is  being  followed 
here,  treats  L.  planetica  as  a  variety  of  L.  erratiea  having  a  more  well- 
developed  thallus  than  the  variety  erratiea.  Lowe  regards  L.  micytho  as 
a  yellowish  form  of  var.  planetica.  The  separations  of  L.  erratiea  and 
L.  cyrtidia  have  already  been  discussed  under  the  latter  species.  L.  sylvi¬ 
cola  differs  from  L.  erratiea  in  having  a  greenish  or  greenish  black 
hypothecium  with  a  doubtfully  distinguished  pale  bluish  black  exciple 
(Lowe,  1939)  as  opposed  to  a  reddish  brown  to  almost  black  hypo¬ 
thecium  and  an  exciple  greenish  black  externally  and  hyaline  within. 

This  common  species  is  found  on  pebbles  in  exposed  fields  and 
downs,  and  is  particularly  abundant  in  well-lighted  areas  on  the  Ronkon- 
koma  moraine  (figure  60).  Some  observations  on  its  development  have 
been  presented  on  p.  43. 

Distribution  —  Eastern  United  States  west  to  Minnesota  (Lowe, 
1939):  Temperate  element.  East  Temperate  subelement;  Europe  (ibid). 

54.  Lecidea  granulosa  (Ehrh.)  Ach.  Meth.  Lich.  65.  1803.  Lichen 
granulosa  Ehrh.  PI.  Crypt.  Exs.  145.  1785. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25644  (64), 
25656  (64),  25788  (86);  Brodo  655  (79),  1900  (114),  1939  (85), 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  1  69 

3372  (94),  3401  (75),  1404  (83);  Shinnecock  Hills,  Latham  7873, 
February  14,  1938  (Latham);  Southold,  Latham  7863,  February  11,  1938 
(Latham);  North  Sea,  Latham  28128,  May  16,  1955  (Latham);  River- 
head,  Peck,  September  (NYS). 

Lecidea  granulosa  is  similar  in  some  respects  to  L.  aeruginosa,  but 
the  two  are  quite  distinct  on  Long  Island  (see  discussion  under  L.  aeru¬ 
ginosa).  It  is  known  to  grow  on  old  wood  as  well  as  soil  but  is  restricted 
on  the  island  to  sandy  soil. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  North  Carolina, 
Michigan,  Minnesota,  Arizona,  Black  Hills,  Washington,  Alaska,  Sas¬ 
katchewan;  Northern  United  States  (Fink,  1935):  Temperate  element, 
North  Temperate  subelement;  Europe;  Asia  (Vainio,  1928). 

55.  Lecidea  macrocarpa  (DC.  in  Lam.  &  DC.)  Steud.  Nomencl.  Bot. 
245.  1824.  Patellaria  macrocarpa  DC.  in  Lam.  &  DC.  FI.  Franc,  ed.  3 
2:  347.  1805. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25592  (52). 

There  has  been  much  disagreement  concerning  the  name  of  this 
species.  Fink  (1935)  discussed  it  under  the  name  L.  platycarpa  Ach.,  and 
Lowe  (1939)  following  Vainio  (1909,  1934)  used  L.  steriza  (Ach.) 
Vain.  Clauzade  and  Rondon  (1959)  recently  considered  the  species  under 
the  name  Lecidea  contigua  (Hoffm.)  Th.  Fr.  Most  other  workers  have 
used  L.  macrocarpa. 

The  epithet  “ macrocarpa ”  was  first  used  at  the  species  level  in  the 
genus  Patellaria  by  DeCandolle  in  1805,  which  makes  it  the  oldest  avail¬ 
able  name.  “Steriza"  was  only  considered  at  the  infraspecific  level  (L.  con- 
fluens  8  L.  steriza  Ach.)  until  Vainio  raised  it  to  a  species  in  1909. 
Lecidea  platycarpa  was  not  described  until  1810  by  Acharius.  Vainio 
(1934)  states  that  Theodor  Fries  used  the  name  contigua  incorrectly  in 
referring  Hoffman's  Verrucaria  contigua  to  the  genus  Lecidea.  Fries’  lichen 
was  L.  macrocarpa  but  Hoffman’s  name  referred  to  a  different  species. 

The  Long  Island  specimen  has  a  rather  well-developed  continuous 
to  cracked  and  areolate  thallus.  It  was  found  on  a  siliceous  roadside 
pebble. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Tennessee,  Michi¬ 
gan,  Minnesota,  Idaho,  Alaska,  Saskatchewan,  Baffin  Island:  Arctic- 
boreal  element;  circumboreal. 

56.  Lecidea  myriocarpoides  Nyl.  Flora  48:  355.  1865. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2535  (49),  2362 
(42),  3880  (62),  3892  (112). 

This  species  was  found  only  on  well-illuminated,  hard  lignum. 

Distribution  —  Eastern  United  States  and  California  (Fink,  1935, 
Lowe,  1  939  );  Europe. 

57.  Lecidea  nylanderi  (Anzi)  Th.  Fr.  Lich.  Scand.  1:  462.  1874. 
Biatora  nylanderi  Anzi.  Cat.  Lich.  Sondr.  75.  1860. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1400  (65),  1953 
(85),  2000  (51),  2549A  (73). 


170  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  very  small,  reddish  brown  apothecia  and  the  subglobose  to 
globose  spores  of  this  species  easily  distinguish  it  from  other  pine  bark 
lichens.  On  Long  Island  it  is  limited  to  the  bark  of  Finns  rigida.  Culber¬ 
son  (1958a),  studying  the  pine-inhabiting  lichen  vegetation  of  North 
Carolina,  found  Lecidea  nylanderi  only  in  the  mountains  of  North  Caro¬ 
lina.  Finns  rigida  is  also  found  only  in  the  mountains.  This  correlation 
may  indicate  a  very  high  degree  of  substrate  specificity,  but  since  the 
specificity  of  the  species  was  not  indicated  in  that  paper,  and  since  other 
pines  occur  in  the  mountains,  no  such  conclusion  can  be  made.  The 
species  is  found  on  Finns  ponderosa  in  the  Black  Hills  of  South  Dakota 
and  was  also  collected  twice  ( Brodo  4122,  4489 )  on  Finns  rigida  in  the 
Cape  Cod  region  of  Massachusetts. 

Distribution  —  Adirondack  Mountains  of  New  York,  Massachusetts, 
California,  (Lowe,  1 939) ;  North  Carolina,  Wisconsin,  Black  Hills,  Mani¬ 
toba:  Temperate  element,  North  Temperate  subelement;  Europe;  Asia 
(Vainio,  1928). 

58.  Lecidea  scalaris  ( Ach.)  Ach.  Meth.  Lich.  78.  1803.  Lichen 
scalaris  Ach.  Kgl.  Vet.  Akad.  Nya  Hand!.  127,  tab.  5,  f.  2.  1795. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3508  (10).  SUFFOLK 
COUNTY:  19  specimens  collected  by  Brodo  and/or  Imshaug. 

As  with  L.  anthrocophila,  this  species  is  treated  under  Psora  by  Fink 
(1935). 

Lecidea  scalaris  has  a  high  specificity  for  the  bark  of  Finns  rigida 
but  is  not  restricted  to  it  (figure  43).  Barkman  (1958,  p.  38)  and  Lowe 
(1939)  state  that  the  species  is  commonly  found  on  burned  wood,  and 
this  is  certainly  true  on  Long  Island  where  it  is  often  found  on  charred 
pine  bark  (see  p.  45).  Acer  saccharinnm  and  Acer  rubrum  were  the 
preferred  substrates  in  an  area  in  central  New  York  (Brodo,  1959).  The 
reasons  for  these  preferences  are  not  clear,  although  all  the  substrates  are 
highly  acid.  Ochsner  (1928  in  Barkman,  1958  p.  102)  stated  that 
L.  scalaris  is  nitrophobus,  but  this  is  yet  to  be  proven. 

Distribution  —  Central  New  York,  North  Carolina,  Arizona,  Black 
Hills,  Washington,  Saskatchewan:  Temperate  element,  North  Temper¬ 
ate  subelement  (?);  Europe;  Asia  (Lowe,  1939). 

59.  Lecidea  uliginosa  (Schrad.)  Ach.  Meth.  Lich.  43.  1803.  Lichen 
nliginosus  Schrad.  Spic.  FI.  Germ.  1:  88.  1794. 

Material  seen  —  SUFFOLK  COUNTY:  22  specimens  collected  by 
Imshaug  and/or  Brodo. 

Laundon  (1960)  discusses  in  detail  the  similarities  of  this  species 
with  Lecidea  oligotropha  Laund.  The  latter  is  mainly  characterized  by 
its  coarsely  granulose  to  verruculose,  pale  brown  to  yellowish  thallus.  In 
contrast,  L.  uliginosa  has  a  finely  granular  to  almost  leprose  dark  brown 
to  black  thallus.  Only  one  North  American  specimen  of  L.  oligotropha 
(from  Minnesota)  is  cited  by  Laundon. 

Lecidea  uliginosa  often  forms  conspicuous  tar-like  patches  on  partially 
stabilized  sand.  Closer  examination  will  reveal  tiny  black  apothecia  scat- 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


171 


tered  among  the  dark  brown  thalline  granules.  Alvin  (1960)  reported  the 
species  as  occurring  in  dune  communities  in  southern  England,  especi¬ 
ally  in  the  heath,  ecologically  very  similar  to  some  Long  Island  habitats. 

Distribution  —  Nova  Scotia,  Connecticut,  Michigan,  Indiana,  Min¬ 
nesota,  Black  Hills;  throughout  United  States  (Fink,  1935);  Temperate 
element,  North  Temperate  subelement( ?) ;  Europe;  Asia  (Vainio,  1928). 

60.  Lecidea  varians  Ach.  Syn.  Meth.  Lich.  38.  1814. 

Material  seen  —  SUFFOLK  COUNTY:  37  specimens  collected  by 
Imshaug  and/or  Brodo;  East  Marion,  Latham  11  (22249),  May  3,  1914 
(Latham);  Greenport,  Latham  3984,  April  1,  1927  (Latham);  Orient, 
Latham  70,  May  30,  1914  (Latham). 

This  species  occurs  on  the  bark  of  various  trees  from  completely 
exposed  dune  areas  to  protected  oak  forests  (figure  50). 

Distribution  —  Nova  Scotia,  Connecticut,  Michigan,  Minnesota, 
Washington,  Manitoba;  throughout  the  United  States  (Fink,  1935); 
France  (Acharius,  1814). 

61.  Lecidea  vernalis  (L.)  Ach.  Meth.  Lich.  68.  1803.  Lichen  vernalis 
L.  Syst.  Nat.  3:  234.  1768. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25752  (132), 
Brodo  816  (55) ,  851  (47),  2649  (61);  Greenport,  Latham  1998  (22247), 
February  27,  1927  (Latham);  Greenport,  Latham  22254,  May  14,  1914 
(Latham);  Greenport,  Latham,  March  1,  1923  (Latham). 

This  species  is  distinguished  by  its  strongly  convex,  pale  apothecia 
and  its  fusiform,  occasionally  one-septate  spores.  It  is  not  common  on 
Long  Island,  but  where  it  occurs,  it  often  covers  large  portions  of  the 
tree  trunk.  It  is  found  on  the  bark  of  various  trees,  particularly  in  rain 
tracks  or  in  other  equally  moist  or  humid  microhabitats.  Outside  of  Long 
Island,  the  species  is  known  to  occur  over  moss. 

Distribution  —  Nova  Scotia,  Connecticut,  Michigan,  Wisconsin, 
Minnesota,  Arizona,  Alaska,  Manitoba,  Baffin  Island:  Arctic-boreal  ele¬ 
ment;  circumboreal. 

62.  Lecidea  viridcscens  (Schrad.  in  Gmel.)  Ach.  Meth.  Lich.  62. 
1803.  Lichen  viridescens  Schrad.  in  Gmel.  Syst.  Nat.  2(2):  1361.  1791. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3016(11). 

This  rare  species  was  found  growing  over  rotting  wood.  It  some¬ 
what  resembles  a  Lepraria  in  its  granulose,  effuse  thallus. 

Distribution  —  Michigan,  Minnesota,  Arizona,  Alaska;  Eastern 
United  States  and  California  (Fink,  1935):  Temperate  element,  North 
Temperate  subelement;  Europe;  Asia  (Vainio,  1928). 

29.  CATILLARIA  (Ach.)  Th.  Fr. 

63.  Catillaria  glauconigrans  (Tuck.)  Hasse,  Bryol.  12:102.  1909. 
Biatora  glauconigrans  Tuck.  Proc.  Amer.  Acad.  Arts.  Sci.  12:  179.  1877. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-272  (53), 
2549B  (73). 

This  species,  rare  on  Long  Island,  was  found  only  on  pine  bark. 
Nearing  (1947)  stated  that  it  is  an  oak-  and  pine-dwelling  lichen.  It  is 


172  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

therefore  surprising  and  noteworthy  that  Thomson  (1951)  reported  the 
species  as  growing  on  the  bark  of  aspens  ( Populus  tremuloides) .  This  sub¬ 
strate  usually  bears  a  typically  neutrophytie  community  of  Caloplaca  spp., 
Physcia  spp.,  and  Xanthoria  spp.,  quite  opposite  from  the  communities 
on  highly  acid  conifer  bark. 

Distribution  —  Massachusetts  and  California  (Fink,  1935),  New 
York  (Nearing,  1947),  Michigan,  Arizona,  Manitoba:  Temperate  ele¬ 
ment,  North  Temperate  subelement;  endemic. 

30.  BACIDIA  De  Not. 

64.  Bacidici  atrogrisea  (Del.  in  Hepp)  Korb.  Parerg.  Lich.  133. 
1860.  Biatora  atrogrisea  Del.  in  Hepp,  Flecht.  Europ.  26.  1853. 

Material  seen  —  SUFFOLK  COUNTY :  Orient,  Latham  84B,  May 
10,  1914  (Latham);  Orient,  Latham  791,  April  5,  1914  (Latham); 
Orient,  Latham,  May  6,  1915,  (Latham);  Orient,  Latham ,  April  10,  1921 
(Latham). 

Erichsen  (1957)  listed  B.  atrogrisea  as  a  form  of  B.  endoleuca  (Nyl.) 
Kickx.  Accepting  this  synonomy,  atrogrisea  must  be  regarded  as  the 
proper  name  for  the  species  since  it  is  older  on  the  species  level.  Nylander 
was  the  first  to  use  endoleuca  as  a  species  ( Lecidea  endoleuca  Nyl.  Acta 
Soc.  Sci.  Fenn.  7:460.  1863.) 

Bacidia  hiteola  (Schrad.)  Mudd  (syn.  B.  rubella  [Hoffm.]  Mass.) 
and  B.  fuscorubella  (Hoffm.)  Bausch.  can  sometimes  be  confused  with 
B.  atrogrisea.  Bacidia  hiteola  is  distinguished  by  its  uniformly  pale  to  dark 
brown  or  reddish  apothecia  with  no  pruina,  often  becoming  very  convex  to 
hemispherical.  Bacidia  fuscorubella  differs  in  having  red-brown  to  almost 
black  apothecia,  often  with  conspicuous  white  pruinose  margins.  Both 
these  species  have  thicker  thalli  than  B.  atrogrisea.  The  Long  Island 
material  agrees  very  well  with  the  Migula  exsiccat,  Krypt.  Germ.  no.  52. 

The  species  is  usually  found  on  the  bark  of  various  coniferous  trees. 
Thomson  (1951)  reported  it  from  Michigan  on  Thuja  bark. 

Distribution  —  Connecticut,  Michigan;  Eastern  United  States  (Fink, 
1935):  Temperate  element,  East  Temperate  subelement;  Europe;  Asia 
(Ikoma,  1957). 

65.  Bacidia  chlorantha  (Tuck.)  Fink,  Cont.  U.  S.  Nat.  Herb.  14: 
91.  1910.  Biatora  chlorantha  Tuck.  Proc.  Amer.  Acad.  Arts  Sci.  1:252. 
1847.  (Syn.  Lich.  New  Engl.  60.  1848.) 

Material  seen  —  NASSAU  COUNTY:  Brodo  569  (11).  SUFFOLK 
COUNTY:  Brodo  2001  (51),  2394  (113),  2647  (61),  3437  (sterile) 
( 134),  3446  (134),  3831  (sterile)  (66). 

Lamb  ( 1954)  presented  a  description  and  a  discussion  of  this  species 
and  Thomson  (1951)  compared  it  with  B.  chlorococca,  with  which  it  is 
sometimes  confused.  The  Long  Island  material  agrees  well  with  Lamb's 
description  of  the  specimens  from  Nova  Scotia. 

This  species  is  often  found  sterile,  but  with  many  clusters  of  minute 
brown  pycnidia  containing  pycnoconidia  measuring  1.2  x  0.5  p,.  It  is 
found  on  the  bark  of  various  species  of  deciduous  and  coniferous  trees. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  173 

Distribution  —  Nova  Scotia,  Connecticut,  Smoky  Mountains  of 
North  Carolina  and  Tennessee,  Michigan;  New  England,  New  York, 
Ohio,  Illinois,  Minnesota  (Fink,  1935):  Temperate  element,  Appalachian 
subelement,  Appalachian-Great  Lakes  unit;  endemic. 

66.  Bacidia  chlorococea  (Graewe  in  Stizenb.)  Lett.  Hedw.  52:  131. 
1912.  Lecidea  chlorococca  Graewe  in  Stizenb.  Nova  Acta  Acad.  Leop. 
Carol.  34  (2):  24.  1867. 

Material  seen  —  NASSAU  COUNTY:  Brodo  536  (16),  555  (12), 
568  (11),  1308  (15).  SUFFOLK  COUNTY:  75  specimens  collected  by 
Imshaug  and/or  Brodo;  Riverhead,  Latham,  May  1,  1960  (Latham). 

Degelius  (1940)  described  the  spores  of  his  material  from  Maine 
as  slightly  smaller  than  those  of  typical  European  specimens,  although  the 
specimens  from  Long  Island  fit  the  spore  size  of  the  European  material 
well. 

The  species  is  found  on  a  variety  of  substrates,  including  twigs  and 
bark  of  coniferous  and  deciduous  trees  as  well  as  old  wood.  It  is  found 
in  exposed  and  shaded  localities. 

Distribution  —  Maine,  central  New  York,  North  Carolina,  Michi¬ 
gan,  Wisconsin:  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit;  Europe. 

67.  Bacidia  chlorosticta  (Tuck.)  Schneid.  Guide  Study  Lich.  109. 
1898.  Lecidea  chlorosticta  Tuck.  Proc.  Amer.  Acad.  Arts  Sci.  5:  419. 
1862. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2097  (78);  River- 
head,  Latham  2380,  June  24,  1924  (Latham). 

This  rare  species  is  distinctive  in  many  ways.  Its  paraphyses  appear 
to  be  branched,  giving  the  apothecium  an  ascolocular  appearance,  but  the 
olivaceous,  minutely  verruculose  to  subgranulose  thallus,  the  large-celled 
Trebouxioid  phycobiont,  and  the  lack  of  thick-walled  asci  all  are  char¬ 
acteristic  of  Bacidia  and  not  Micarea.  The  apothecia  are  small,  lead  black, 
and  convex,  with  the  margin  disappearing.  They  are  sessile  or  buried  in 
the  granular  crust,  or  sometimes  they  become  more  or  less  stipitate. 
The  hypothecium  is  dark  brown,  becoming  sordid  blackish  violet  below 
and  olivaceous  above  in  KOH.  The  margins  are  reddish  violet  in  KOH. 

The  species  is  apparently  restricted  to  Chamaecyparis  thyoides,  at 
least  in  the  coastal  plain  region.  Three  specimens  ( Brodo  3676,  3765, 
3768 )  were  collected  in  southern  New  Jersey  on  white  cedar  in  cedar 
bogs  just  as  they  were  on  Long  Island. 

Distribution  —  Connecticut;  Massachusetts,  South  Carolina,  Illinois 
(Fink,  1935);  New  Jersey  (cf.  above):  Temperate  element,  Coastal  Plain 
subelement;  endemic. 

68.  Bacidia  intermedia  ( Flepp  in  Stizenb.)  Arn.  Flora  54:  54.  1871. 
non  Hampe  in  Mass.  Biatora  anomala  var.  intermedia  Hepp  in  Stizenb. 
Nova  Acta  Acad.  Leopold.  —  Carolin.  30  (3):  42.  1863. 

Material  seen  -  SUFFOLK  COUNTY:  Brodo  3209  (33);  Orient. 
Latham  84,  May  10,  1914  (Latham)- 


174 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


It  seems  dear  from  published  descriptions  that  the  Long  Island 
material  te'ongs  to  what  Tuckerman  (1888,  sub  Biatora ),  Fink  (1935), 
and  Erichsen  (  1957)  have  called  Bacidia  effusa  (Sm.  in  Sm.  &  Sowerby) 
Trev.  However,  there  are  a  number  of  problems  involved  in  the  use  of  the 
name  B.  effusa.  First,  the  epithet  effusa  cannot  be  used  for  any  Bacidia 
since  its  basionym,  Lichen  effusus  Sm.  in  Sm.  &  Sowerby  (1808),  is  a  later 
homonym  of  Lichen  effusus  Ach.  ( 1798),  a  synonym  of  Lecanora  saligna 
(Schrad.)  Zahlbr.  (see  Recommendation  72A,  Lanjouw,  1961  ).  Secondly, 
Lichen  effusus  Sm.  in  Sm.  &  Sowerby  is  listed  as  a  synonym  of  Bacidia 
arceutina  (Ach.)  “Arn.”  by  Vainio  (1922).  The  latter  species  as  de¬ 
scribed  by  Vainio  differ  in  many  respects  from  the  Long  Island  specimens. 
These  Long  Island  specimens  do  fit  Vainio’s  description  of  Bacidia  inter¬ 
media  (“Hepp”)  Arn.  and  they  agree  almost  perfectly  with  the  Raben- 
horst  exsiccat  no.  509  (distributed  as  Bacidia  effusa)  cited  by  Vainio 
as  typical  B.  intermedia.  Still  another  name  which  must  be  considered  is 
Bacidia  albescens  (Hepp)  Zwack.,  which  some  authors  (e.g.,  Arnold, 
1884)  considered  as  including  intermedia  as  only  a  form  with  flatter,  larger 
apothecia.  Erichsen  (1957)  used  all  four  names  (B.  effusa,  B.  arceutina, 
B.  intermedia  and  B.  albescens)  as  separate  species  distinguishing  them 
as  follows: 

1.  Apothecia  at  first  light,  darker  in  age,  never  black;  spores  mostly 


20-50  ij.  long . 2 

1.  Apothecia  soon  or  from  the  beginning  dark  to  black;  spores  mostly 

40-60  p.  long  (but  f.  brevispora  is  25-39  \j.  long) . B.  arceutina 

2.  Apothecia  whitish,  flesh  colored,  or  rose . . . 3 

2.  Apothecia  light  brown,  brick  red,  darker  in  age . B.  effusa 

3.  Hymenium  50-60;j.  high;  spores  34-48y.  long1-;  apothecia  remaining 

flat.  0-3-0. 4(0. 6)  mm  in  diameter . B.  intermedia 

3.  Hymenium  35-50  p  high;  spores  20-33  ;j.  long;  apothecia  first  flat,  then 
soon  convex  and  marginless,  0. 2-0.4  mm  in  diameter.  .  .  B.  albescens 
Fink  (1935)  did  not  use  B.  intermedia.  He  separated  B.  arceutina, 
B.  effusa  and  B.  albescens  as  follows: 

1 .  Spores  rarely  more  than  40  p  in  length . 2 


1.  Spores  rarely  less  than  40  p  in  length  (35-50  p).  Hypothecium  yellow¬ 
ish;  disks  pale  light  brown  to  blackish . B.  arceutina 

2.  Hypothecium  hyaline;  apothecia  not  more  than  0.5  mm  across;  disk 

light  pink  to  pale  reddish . B.  albescens 

2.  Hypothecium  pale  yellowish;  apothecia  small,  0.4-0. 8  mm  across; 

disk  pale  flesh  colored  to  reddish  brown . B.  effusa 

It  appears  that  Bacidia  effusa  sensu  Fink  and  Erichsen  is  probably 
synonymous  with  Vainio’s  B.  intermedia,  and  intermedia  is  in  all  likeli¬ 
hood  merely  a  form  of  B.  albescens.  Since  I  have  not  yet  seen  any  authen¬ 
tic  material  of  B.  albescens,  Vainio’s  interpretation  is  followed  at  this 
time.  Unfortunately,  the  epithet  intermedia  on  the  species  level  (Arnold, 
1871)  is  preempted  by  Bacidia  intermedia  Hampe  in  Mass.  (1861),  and 


12  Based  on  Vainio  (1922),  which  in  turn  is  based  on  a  single  specimen. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  175 

therefore  is  invalid.  Until  further  studies  are  done  to  determine  the  cor¬ 
rect  name  for  this  taxon,  however,  B.  intermedia  (Hepp  in  Stizenb.) 
Arn.  will  be  used. 

Distribution  - —  Massachusetts,  Iowa,  California  (Fink,  1935,  sub 
B.  effusa);  Europe;  Asia  (Vainio,  1928,  sub  B.  intermedia) . 

69.  Bacidia  inundata  (Fr.)  Korb.  Syst.  Lich.  Germ.  187.  1855.  Bia- 
tora  inundata  Fr.  Kgl.  Vet.  Akad.  Nya  Flandl.  270.  1822. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  761  (67),  3917  (54). 

This  species  is  usually  found  on  siliceous  rocks  in  or  near  a  stream 
or  brooks  (Hale,  1950;  Thomson,  1951),  but  the  Long  Island  specimens 
were  collected  in  comparatively  dry  habitats  on  concrete.  One  (no.  761) 
was  growing  on  a  concrete  foundation  within  a  few  feet  of  a  swampy 
brook,  and  the  other  was  collected  in  a  shaded  oak  woods,  on  an  old 
concrete  foundation.  However,  Sandstede  (1913)  reported  B.  inundata 
from  brick  walls  and  Fink  ( 1902)  listed  the  species  from  limestone  bluffs 
in  Minnesota.  Tuckerman  (1888)  stated  that  B.  inundata  is  found  “on 
various  rocks,  especially  such  as  contain  lime;  as  also  on  brick;  .  .  .” 

Distribution  —  Connecticut,  Michigan,  Minnesota,  Indiana,  Black 
Hills;  East  of  Rocky  Mountains  (Fink,  1935):  Temperate  element.  East 
Temperate  subelement;  Europe. 

70.  Bacidia  schweinitzii  (Tuck,  in  W.  Dari.)  Schneid.  Guide  Study 
Lich.  110.  1898.  Biatora  schweinitzii  Tuck,  in  W.  Dari.  FI.  Cestr.  ed.  3. 
447.  1853. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2121  (102),  2757 
(102);  2802  (102). 

This  species  is  unique  among  the  Bacidiae  in  having  Trentepohlia  as 
a  phycobiont  rather  than  Trebouxia.  Lamb  (1954)  discusses  this  fact  and 
some  other  aspects  of  the  history  of  the  species. 

Bacidia  schweinitzii  was  found  in  only  one  locality,  as  a  member  of 
the  Acer  rubrum- bog  community.  It  was  also  found  in  southern  New 
Jersey  (Burlington  County,  Atsion,  Brodo  3558)  on  a  roadside  oak  close 
to  a  bog. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Tennessee,  North 
Carolina,  Oklahoma,  Michigan,  Indiana,  Minnesota;  eastern  United 
States  (Fink.  1935):  Temperate  element,  East  Temperate  subelement; 
endemic. 

71.  Bacidia  cfr.  trisepta  (Naegeli  in  Mull.  Arg.)  Zahlbr.  in  Engler- 
Prantl,  Nat.  Pflanzenf.  1(1):  135.  1907.  Lecidea  trisepta  Naegeli  in  Mull. 
Arg.  Mem.  Soc.  Phys.  Hist.  Nat.  Geneve  16:  403.  1862. 

Material  seen  — -  SUFFOLK  COUNTY:  Brodo  2337  (44);  Green- 
port,  Latham  38166 A,  July  14,  1963  (Latham);  Greenport,  Latham 
38196A,  April  15,  1965  (Latham). 

Bacidia  trisepta,  except  for  f.  saxicola  (Korb.)  Lettau,  is  mainly 
known  from  lignum  and  bark.  The  ascocarps  appeared  to  be  ascolocular  in 
the  Long  Island  specimen.  Since  I  have  not  examined  the  type  and  the 


176  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Long  Island  material  was  questionable,  I  will  not  transfer  the  species  into 
Micarea  where  it  might  very  well  belong. 

The  specimens  were  all  collected  on  shaded  granitic  rock. 

Distribution  —  Massachusetts  (Fink,  1935):  Black  Hills;  Europe. 

72.  Bacidia  umbrina  (Ach.)  Bausch,  Verh.  Nat.  Ver.  Carls.  4:103. 
1869.  Lecidea  umbrina  Ach.  Lich.  Univ.  183.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2738  (111). 

The  distinctive  twisted  and  curved  spores  of  this  species  easily  sepa¬ 
rate  it  from  all  other  Bacidiae  on  Long  Island.  Accurate  measurements  of 
the  spore  length  were  difficult  due  to  the  strong  curvature  of  the  spores, 
and  the  values  appear  to  he  somewhat  lower  than  those  reported  by  Hill¬ 
man  &  Grummann  (1957)  or  Erichsen  (1957)  ( 1 5-20  rather  than 
1 7-40  [j.  in  length) . 

The  ecology  of  the  specimen  found  on  Long  Island  was  extremely 
unusual  for  the  species.  It  was  found  in  the  hygrohaline  stratum  on  a 
granite  boulder  above  the  littoral  zone  but  well  within  the  storm  tidal 
level,  and  certainly  exposed  to  salt  spray  in  windy  weather.  Growing 
alongside  the  specimen  was  Acarospora  fuscata  (p.  61). 

Distribution  —  Maine,  Connecticut,  Tennessee,  Minnesota;  northern 
United  States  (Fink,  1935):  Temperate  element,  North  Temperate  sub¬ 
element;  Europe. 

31.  RHIZOCARPON  Ram. 

73.  Rhizocarpon  cinereovirens  ( Miill.  Arg.)  Vain.  Acta  Soc.  Faun. 
FI.  Fenn.  53  (1):  336.  1922.  Patellaria  cinereovirens  Miill.  Arg.  Flora 
51:  49.  1868. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2173  (99),  3265 
( 1 19),  3899  (112). 

The  very  lightly  tinted  or  hyaline  1 -septate  spores  of  this  species 
give  it  the  appearance  of  a  saxicolous  Catillaria  or  a  light-spored  Buellia 
(especially  B.  stigmaea).  However,  gelatinous  episporic  sheaths  are  usu¬ 
ally  conspicuous,  indicating  its  true  position. 

Runemark  (1956)  identified  both  norstictic  and  stictic  acids  from 
R.  cinereovirens  by  chromatographic  analysis.  The  presence  of  norstictic 
acid  in  the  medulla,  an  unusual  feature  among  the  Catocarpons,  was  de¬ 
tected  in  two  of  the  Long  Island  specimens.  The  third  specimen  (Brodo 
3899)  was  KOH  +  yellow;  chromatography  showed  the  presence  of 
stictic  acid,  but  not  norstictic  acid.  Unfortunately,  the  Long  Island  ma¬ 
terial  was  too  scanty  to  enable  a  more  thorough  chemical  analysis.  How¬ 
ever,  the  presence  of  stictic  and  norstictic  acids  together  is  by  no  means 
uncommon  (cfr.  Parmelia  conspersa,  P.  Iiypotropa,  etc.). 

Distribution  —  Black  Hills;  Minnesota  (Fink,  1935);  Europe. 

74.  Rhizocarpon  grande  (Florke  in  Flot.)  Arn.  Flora  54:  149.  1871. 
Lecidea  petraea  var.  fuscoatra  f.  grandis  Florke  in  Flot.  Flora  1 1 :  690. 
1828. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3850  (76). 

Rhizocarpon  grande  was  discussed  at  length  by  Degelius  (1940, 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


177 


1941).  Degelius  (1940)  mentioned  the  KOH  +  yellow  to  testaceous  re¬ 
action  of  the  medulla  as  well  as  the  C  +  red  reaction.  The  substances 
responsible  for  these  reactions  were  identified  by  Runemark  (1956)  as 
stictic  and  gyrophoric  acids.  Stictic  acid  was  found  in  the  Long  Island 
specimen  (paper  chromatography),  and  the  C  +  red  reaction  indicates 
that  gyrophoric  acid  is  probably  present  as  well. 

The  specimen  was  found  on  an  exposed  granite  boulder. 

Distribution  —  Maine,  Tennessee,  Michigan,  Minnesota,  Idaho, 
Black  Hills,  Washington,  Saskatchewan,  Manitoba,  Baffin  Island:  Arctic- 
boreal  element;  circumboreal. 

75.  Rhizocarpon  intermedium  Degel.  Ark.  Bot.  30A  (3):  43.  1941. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1903  (114),  2662a 

(108),  3271  (119). 

The  Long  Island  specimen  agreed  perfectly  with  the  type  material 
(US).  The  type  specimen  contained  stictic  acid  (by  chromatography),  as 
did  the  Long  Island  material  (except  one  poorly-developed  specimen). 
I  also  collected  the  species  on  Cape  Cod  (Massachusetts)  ( Brodo  3947, 
4201,  4207a). 

Distribution  —  Tennessee,  Massachusetts  (see  above);  endemic. 

76.  Rhizocarpon  obscuratum  (Ach.)  Mass.  Ricerch.  Auton.  Lich. 
103.  1852.  Lecidea  petraea  y  L.  obscurata  Ach.  Lich.  Univ.  156.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25599  (52), 
Brodo  946  (53),  1753  (126),  1967  (91),  2340  (44),  2719a  (111), 
2740  (111),  3078c  (128),  3285  (119),  3902  (112);  Orient,  Latham 
7413,  May  1,  1933  (Latham);  Quogue,  Latham  28254B  (Latham); 
Shinnecock,  Latham  27288,  May  8,  1945  (Latham). 

This  species  is  apparently  extremely  variable,  with  many  forms  hav¬ 
ing  been  described  for  it  (Erichsen  [1957]  included  ten).  Of  the  many 
forms,  f.  reduction  (Th.  Fr. )  Eitn.  seems  to  be  most  common  on  Long 
Island.  This  form  is  distinguished  by  a  "more  granulose  thallus,  smaller 
apothecia  with  thinner  and  disappearing  margin,  indistinctly  papillated 
disk  and  submurale  (not  morale)  spores”  (Degelius,  1940).  All  Long 
Island  specimens  lacked  any  clearly  positive  chemical  tests,  although 
Runemark  ( 1956)  reported  both  stictic  and  gyrophoric  acids  from  a  speci¬ 
men  which  he  tested. 

Rhizocarpon  orphninum  (Vain.)  Zahlbr.  is  very  similar  to  R.  obscu¬ 
ratum  but  differs  in  having  a  KOH  +  violet  or  magenta  reaction  in  the 
exciple  and  epithecium  (Laundon,  1960). 

Rhizocarpon  obscuratum  is  common  on  pebbles  and  small  stones  and 
is  often  associated  with  Lecidea  erratica. 

Distribution  —  Maine,  Minnesota  (Fink,  1935);  Nova  Scotia,  Con¬ 
necticut,  Tennessee,  Saskatchewan;  Greenland  (Lynge,  1940c);  Europe; 
Asia  (Lynge,  1928) 

77.  Rhizocarpon  plicatile  (Leight.)  A.  L.  Sm.  Monogr.  Brit.  Lich. 
2:  197.  1911.  Lecidea  plicatilis  Leight.  Ann.  Mag.  Nat.  Hist.  IV.  4:  201. 
1869. 


178  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2336  (44),  3076 
(128). 

Rhizocarpon  plicatile  was  found  on  well-illuminated  or  partially 
shaded  boulders.  I  have  collected  specimens  from  the  Adirondack  Moun¬ 
tains  of  New  York. 

Distribution  —  Nova  Scotia,  Maine,  northern  New  York  (see 
above),  North  Carolina;  Europe. 

STEREOCAULACEAE 

32.  PYCNOTHELIA  (Ach.)  Duf. 

78.  Pycnothelia  papillaria  (Ehrh.)  Duf.  Ann.  Gen.  Sci.  Phys.  Brux. 
8:  5.  1817.  Lichen  papillaria  Ehrh.  Phytophyl.  no.  100.  1780. 

Material  seen  - — -  NASSAU  COUNTY:  Brodo  3345  (8);  Plain  Edge, 
S.  Cain  371,  372,  1936,  Andropogonetum  Hemsteadi  (NY).  SUFFOLK 
COUNTY:  Brodo  59-177  (100B),  1177  (101),  1559  (103),  1682  (88), 
1750  (126),  1752  (126),  1980  (91),  2015  (51),  2533  (49),  3005 
(17),  841  (55);  16  specimens  collected  by  Latham  (Latham);  Orient, 
Booth,  August,  1877  (FH);  Orient,  Latham  13,  V.  1914  (FH);  Orient 
Point,  Latham,  1927  (NY);  Montauk  Point,  R.  H.  Torrey,  1933  (NY); 
Selden,  S.  Cain  348,  359,  360,  1936  (NY);  Coram,  R.  H.  Torrey,  1936 
(NY);  Calverton,  R.  H.  Torrey,  1936  (NY);  East  of  Calverton,  R.  H. 
Torrey,  1936  (NY);  Route  112,  north  of  Coram,  R.  H.  Torrey,  1936 
(NY). 

The  important  characters  which  separate  Pycnothelia  from  Cladonia ; 
e.g.  pseudopodetia  rather  than  podetia  (see  Lamb,  1951),  and  septate 
spores  rather  than  nonseptate  spores,  have  for  some  reason  been  ignored 
in  the  recent  past  by  most  workers,  with  the  notable  exception  of  Watson 
(1953)  and  Mattick  (1938).  However,  even  Mattick  (1940)  later  chose 
to  regard  Pycnothelia  as  part  of  Cladonia  “for  practical  reasons.”  Hale 
and  Culberson  ( 1966)  recognized  the  genus  in  their  new  checklist. 

Pycnothelia  papillaria  seems  to  be  narrowly  restricted  to  well- 
illuminated  localities  on  eroding  sandy  loam  (p.  56;  figure  73). 

Distribution  —  Temperate  element.  East  Temperate  subelement: 
Europe  (map:  Sandstede,  1932). 

33.  STEREOCAULON  Hoffm. 

79.  Stereocaulon  saxatile  Magn.  Gdteb.  Kgl.  Vet.  Sanih.  Hand!.  IV. 
30:  41.  1926. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3852  (76);  (locality 
unknown),  Latham  38187  (No.  7),  April  1924  (Latham). 

Lobaric  acid  and  atranorin  were  demonstrated  by  recrystallization 
in  GAW  and  GAoT,  respectively,  in  the  Long  Island  and  Cape  Cod 
specimens.  These  chemical  constituents  were  reported  for  this  species  by 
Lamb  (1951)  and  Ramaut  (1962). 

The  epithet  evolutoides  was  published  as  a  variety  of  S.  paschale  by 
Magnusson  in  1926  and  was  first  used  on  the  species  level  by  Frey  in 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


179 


1932.  It  is  necessary,  therefore,  to  refer  to  this  species  as  S.  saxatile, 
although  most  recent  authors  treat  saxatile  as  a  variety  of  evolutoides. 

A  specimen  of  this  species  in  much  better  condition  than  the  Long 
Island  material  was  found  on  Cape  Cod  (East  Dennis,  Brodo  4467). 
Both  specimens  were  growing  on  granite  boulders,  the  former  in  partial 
shade  and  the  latter  in  full  sun. 

Distribution  —  Nova  Scotia,  Massachusetts,  Ontario,  Saskatchewan: 
Temperate  element,  North  Temperate  subelement(?)  (see  Ahti,  1964); 
Europe.  Listed  as  an  “amphiatlantic,  boreal”  species  by  Lamb  (1951). 

BAEOMYCETACEAE 

34.  BAEOMYCES  Pers. 

80.  Baeomyces  roseus  Pers.  Neue  Ann.  Bot.  1:  19.  1794. 

Material  seen  —  NASSAU  COUNTY:  Brodo  59-114  (12),  2526 

(5).  SUFFOLK  COUNTY:  lmshaug  25560  (52),  25583  (52), 

25688  (72);  Brodo  59-179  (54),  830  (55),  836  (55),  975  (63), 
1274  (31),  1223  (100A),  1686  (88),  1984  (91),  2003  (51 ),  2987  (26), 
3081  (128),  3342  (76);  13  specimens  collected  by  Latham  (Latham); 
Wildwood  State  Park,  S.  Smith  12669,  October  17,  1952  (NYS). 

This  species  is  usually  found  on  eroding  sandy  loam,  especially  on 
the  moraines  (figure  53),  and  is  often  associated  with  Pycnothelia  papil- 
laria. 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  Connecticut, 
Tennessee;  Appalachian-Great  Lakes  distribution  (Hale,  1961a):  Tem¬ 
perate  element,  Appalachian  subelement,  Appalachian-Great  Lakes  sub¬ 
element  (?);  Europe,  Asia  (circumboreal:  Sandstede,  1932). 

CLADONIACEAE 

35.  CLADONIA  Wigg. 

Subgenus  Cladonia 
Section  CLAUSAE  Korb 
Subsection  Cocciferae  Del. 

Series  Subglaucescentes  Vain. 

81.  Cladonia  floerkeana  ( Fr. )  Florke,  Clad.  Comm.  99.  1828. 
Cenomyce  floerkeana  Fr.  Lich.  Suec.  Exs.  82.  1824. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-161  (83),  2076 
(38),  2996  (17),  3426  (134);  Southold,  Latham  7573  (+  7581, 
+  7588,  +  7590),  January  3,  1934  (Latham). 

This  species  is  found  on  the  ground  in  open  sandy  or  grassy  fields. 
Distribution  —  Vermont,18  Massachusetts,  Connecticut,  New  Jersey, 
Tennessee,  Michigan:  Temperate  element.  East  Temperate  subelement(  ?); 
Europe;  Asia. 

13 In  addition  to  those  locality  references  given  on  nage  1 12.  the  following  refer¬ 
ences  pertain  to  Cladonia :  Vermont  (Evans,  1947),  Connecticut  (Evans.  1930. 
1944),  New  Jersey  (Evans,  1935),  Tennessee  (Mozingo,  1961),  Michigan 
(Evans,  unpublished  key  to  the  Cladoniae  of  Michigan).  References  to  pres¬ 
ence  in  Asia  are  based  an  Asahina  (1950)  unless  otherwise  stated. 


180  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

82.  Cladonia  bacillaris  (Ach.)  Nyl.  Bot.  Sallsk.  Faun.  FI.  Fenn. 
Forh.  8:  179.  1866.  Baeomyces  bacillaris  Ach.  Meth.  Lich.  329.  1803. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  G.  B.  Brainerd, 
1860’s?  (BKL  031984).  NASSAU  COUNTY:  Brodo  550A  (12),  1500 
(9),  1506  (14).  SUFFOLK  COUNTY:  112  specimens  collected  by 
Imshaug  and/or  Brodo;  39  specimens  collected  by  Latham  (Latham); 
Orient,  Latham  209,  May  10,  1914  (FH);  Barling  Hollow  (=  Baiting 
Hollow?),  R.  H.  Torrey,  1934  (NY);  Holtsville,  R.  H.  Torrey,  1936 
(NY);  Southold,  R.  H.  Torrey,  1937  (NY);  Wyandanch  Club  Game 
Reserve  south  of  Smithtown,  R.  H.  Torrey,  1937  (NY);  E.  of  Green- 
port,  S.  Smith  17855,  March  13,  1955  (NYS). 

Cladonia  bacillaris  is  one  of  the  most  common  lichens  on  Long 
Island.  The  species  is  very  variable,  having  numerous  sterile  and  fertile 
forms.  Red  apothecia  are  present  on  approximately  50  percent  of  the 
specimens  and  appear  either  as  conspicuous  hemispherical  terminal  caps 
or  mere  dots  of  red  at  the  podetial  summits.  The  podetia  either  taper 
very  gradually  to  a  point,  are  almost  entirely  uniform  in  diameter,  or 
are  distinctly  clavate. 

The  species  is  found  on  a  variety  of  substrates,  including  soil,  tree 
bases,  and  rotten  wood,  but  it  is  found  most  frequently  on  wood. 

Distribution  —  Vermont,  Massachusetts,  Connecticut,  New  Jersey, 
Michigan,  Indiana,  Minnesota,  Oklahoma,  Arizona,  Black  Hills,  Wash¬ 
ington,  Alaska,  Saskatchewan,  Manitoba,  Ontario:  Temperate  element, 
North  Temperate  subelement;  Europe,  Asia. 

83.  Cladonia  macilenta  Hoffm.  Deutschl.  FI.  2:  126.  1796. 

Material  seen  —  SUFFOLK  COUNTY:  16  specimens  collected  by 

Imshaug  and/or  Brodo;  Montauk,  Hither  beach,  Latham  24001,  24023, 
October  28,  1945  (Latham);  Amagansett,  Latham  25991,  March  11, 
1947  (Latham);  Greenport,  Latham  27479,  April  30,  1950  (Latham); 
Flanders,  Latham  24775,  April  8,  1946  (Latham);  Riverhead,  Latham 
33321,  February  6,  1953  (Latham);  East  Marion,  Latham,  September  1, 
1947  (Latham);  Riverhead,  Latham,  May  16,  1960  (Latham);  Orient, 
Latham  215,  May  23,  1914  (FH);  Orient,  Latham  209,  May  10,  1914 
(FH). 

Cladonia  macilenta  closely  resembles  C.  bacillaris,  the  two  being 
best  separated  by  their  chemistry:  C.  macilenta  has  thamnolic  acid  and 
C.  bacillaris  does  not.  Although  C.  macilenta,  like  C.  bacillaris,  is  found 
on  many  different  substrates,  it  is  usually  found  on  sandy  soil. 

Distribution  —  Vermont,  Massachusetts,  Connecticut,  New  Jersey, 
Tennessee,  North  Carolina,  Michigan,  Ontario,  Minnesota,  Black  Hills, 
Washington,  coastal  Alaska:  Temperate  element,  North  Temperate  sub¬ 
element  (?);  Europe;  Asia. 

84.  Cladonia  vulcanica  Zoll.  Natur-et  Geneeekundig  Arch.  Neerl. 
Indie  1:  396.  1847. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25821  (86), 
25826  (86);  Brodo  2142  (102),  2150  (102);  Northwest,  Latham 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


181 


27458,  April  12,  1948  (Latham);  Flanders,  Latham  24762,  April  8, 
1946  (Latham);  Riverhead,  Latham  32871 ,  April  18,  1955  (Latham); 
North  Sea,  Latham  32317,  35349,  March  26,  1954  (Latham);  Riverhead, 
Latham,  May  2,  1957  (Latham);  Riverhead,  Latham,  May  16,  1960 
(Latham). 

The  presence  of  thamnolic  acid  in  C.  vulcanica  distinguishes  this 
species  from  the  very  similar  C.  didyma.  Both  species  are  found  on  rot¬ 
ting  logs  in  bogs  (figure  36).  It  is  interesting  that  C.  vulcanica  was  found 
to  be  abundant  in  the  white  cedar  bogs  of  Cape  Cod,  but  was  never 
collected  in  my  studies  of  similar  bogs  in  southern  New  Jersey.  Exactly 
the  reverse  was  true  of  C.  didyma. 

Distribution  —  South  America,  and  from  New  Jersey  to  Florida 
(Evans,  1952):  Tropical  element,  Coastal  Plain  subelement;  Asia. 

85.  Cladonia  didyma  (Fee)  Vain.  Acta  Soc.  Faun.  FI.  Fenn.  4:  137. 
1887.  Scyphophorous  didymus  Fee,  Essai  Crypt.  Ecorc.  Off.  98  and  101, 
pi.  3,  f.  13,  1824. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2106B  (86),  2132 
(102);  Montauk,  Hither  Beach,  Latham  24018,  October  28,  1945 
(Latham);  Riverhead,  Latham  29580,  August  7,  1950  (Latham);  River¬ 
head,  Latham  ( 36865)1 ,  May  16,  1960  (Latham);  Northwest,  Latham 
26436,  April  10,  1947  (Latham). 

A  discussion  of  some  aspects  of  the  ecology  and  taxonomy  of  this 
species  can  be  found  with  the  comments  on  C.  vulcanica. 

Distribution  —  Connecticut  to  Florida  along  the  coast:  Tropical 
element,  Coastal  Plain  subelement;  much  of  South  America,  Africa, 
Hawaii,  Ceylon,  Japan  (map:  Sandstede,  1932),  but  not  listed  from  Japan 
by  Asahina  (1950). 

Series  Stramineoflavidae  Vain. 

86.  Cladonia  incrassata  Florke,  Clad.  Comm.  21.  1828. 

Material  seen  —  NASSAU  COUNTY:  Brodo  59-113  (12),  562 

(13),  564  (11),  3512  (10).  SUFFOLK  COUNTY:  29  specimens  col¬ 
lected  by  Imshaug  and/or  Brodo;  39  specimens  collected  by  Latham 
(Latham). 

This  species  is  narrowly  restricted  to  rotting  wood  and  to  pine  bases. 
As  in  Cladonia  cristatella,  if  podetia  are  produced,  they  are  always 
capped  by  large  red  apothecia. 

Distribution  —  Along  the  coast  from  Nova  Scotia  to  Florida  (Evans, 
1952):  Temperate  element,  Coastal  Plain  subelement:  Europe;  Asia. 

87.  Cladonia  cristatella  Tuck.  Amer.  Sci.  25:  428.  1858. 

Material  seen  —  KINGS  COUNTY:  Gowanus,  G.  B.  Brainerd, 

(1866?),  on  ground  ( BKL) .  NASSAU  COUNTY:  Brodo  538  (16),  544 
(12),  1305  (15),  1504  (14),  3193  (6),  3347  (8),  3496  (4).  SUFFOLK 
COUNTY:  93  specimens  collected  by  Imshaug  and/or  Brodo;  61  speci¬ 
mens  collected  by  Latham  (Latham);  Orient  Point,  Latham,  November  6, 
1911  (NYS);  near  Orient,  Latham  17,  1914  (FH);  near  Orient,  Latham 
27,  1914  (FH);  ?,  Latham  15,  1914  (FH);  Orient,  Latham  191,  May  20, 


182  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

1914  (FH);  East  of  Calverton,  R.  H.  Torrey,  1936  (NY);  Holtsville, 
R.  H.  Toney,  1937  (NY);  Pikes  Beach,  Westhampton,  R.  H.  Torrey, 
1936  (NY);  Selden,  R.  H.  Torrey,  1936  (NY);  Selden,  S.  Cain  349, 
1936  (NY);  2.3  miles  SW  of  Riverhead,  5.  Smith  11850,  11851, 
11849,  August  14,  1952  (NYS). 

Cladonia  cristatella  is  common  and  widespread  on  Long  Island, 
occurring  on  a  variety  of  substrates  in  a  multitude  of  forms  (p.  110). 

Distribution  —  Eastern  United  States  (Sandstede,  1939) :  Temperate 
element.  East  Temperate  subelement;  endemic. 

88.  Cladonia  deformis  (L.)  Hoffm.  Deutschl.  FI.  2:  120.  1796. 
Lichen  deformis  L.  Sp.  PI.  1152.  1753. 

Material  seen  -  SUFFOLK  COUNTY:  Montauk  Point,  R.  H.  Tor¬ 
rey,  1933  (NY). 

This  species,  found  only  once  on  Long  Island,  is  very  similar  to 
C.  pleurota,  differing  in  having  farinose  soredia  and  podetial  cups  which 
are  often  lacerate  and  have  involute  margins. 

Distribution  —  Vermont,  Massachusetts,  Connecticut,  Michigan, 
Ontario,  Minnesota,  Black  Hills,  Washington,  Alaska,  Saskatchewan, 
Manitoba,  Canadian  East  Arctic:  Arctic-boreal  element;  circumboreal. 

89.  Cladonia  pleurota  (Florke)  Schaer.  Enum.  Crit.  Lich.  Eur.  186. 
1850.  Capitularia  pleurota  Florke,  Mag.  Ges.  naturf.  Freunde,  Berlin 
2:  218.  1808. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3344  (8);  Massa- 
pequa,  S.  Cain  35,  June  20,  1935  (NY).  SUFFOLK  COUNTY:  20  speci¬ 
mens  collected  by  Imshaug  and/or  Brodo;  Riverhead,  Latham  7707, 
May  1,  1937  (Latham). 

Sterile  podetia  of  C.  pleurota  bear  many  resemblances  to  sterile 
C.  chlorophaea,  and  the  two  are  often  found  together  on  various  types 
of  soil.  The  yellow  color  of  C.  pleurota  (due  to  usnic  acid)  distinguishes 
the  two  in  the  field.  In  addition.  Long  Island  material  of  C.  chlorophaea 
almost  always  can  be  shown  to  contain  grayanic  acid  which  is  absent  in 
C.  pleurota.  Its  similarity  to  C.  deformis  has  already  been  mentioned. 

Clador.ia  pleurota  grows  well  on  eroded,  sandy  loam  as  well  as  on 
mossy  soil  and  so  has  a  broad  distribution  over  both  moraines  (figure  56). 
It  is  also  occasionally  found  on  tree  bases. 

Distribution  —  Nova  Scotia,  Vermont,  Massachusetts,  Connecticut, 
New  Jersey,  Michigan,  Ontario,  Black  Hills,  Alaska,  Manitoba,  Canadian 
East  Arctic:  Arctic-boreal  element;  Europe;  Asia;  “hemiarctic”  (Ahti, 
1964). 

Subsection  Ochroleucae  Fr. 

90.  Cladonia  carneola  (Fr. )  Fr.  Lich.  Eur.  233.  1831.  Cenomyce 
carneola  Fr.  Sched.  Crit.  4:  23.  1825. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2693  (110);  Mon¬ 
tauk,  R.  H.  Torrey,  1933  (NY). 

The  presence  of  barbatic  acid  and  farinose  rather  than  granulose 
soredia  are  usually  sufficient  to  separate  sterile  specimens  of  this  rare 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  183 

species  from  the  more  common  C.  pleurota.  Fertile  material  is  easily 
distinguished,  since  the  apothecia  are  brown  rather  than  red. 

Cladonia  carneola  is  found  on  well-illuminated  eroding  soil. 

Distribution  - —  Black  Hills,  Washington,  coastal  Alaska,  British 
Columbia,  Saskatchewan,  Manitoba:  Arctic-boreal  element;  circumboreal 
(Sandstede,  1939;  Ahti,  1964). 

91.  Cladonia  piedmontensis  Merr.  Bryologist  27:  22.  1924. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3352  (8).  SUFFOLK 

COUNTY:  Imshaug  25611  (116),  Brodo  2821  (115);  Montauk  Point, 
R.  H.  Torrey,  1933  (NY). 

Cladonia  substraminea  Nyl.  (p.p.)  is  listed  as  a  synonym  of  C. 
piedmontensis  by  Fink  (1935).  It  is  also,  in  part,  a  synonym  of  C.  crista- 
tella  f.  ochrocarpia  Tuck.  (Evans,  1930;  Fink,  1935).  Until  the  type  is 
examined  and  the  true  identity  of  C.  substraminea  is  determined,  the 
name  C.  piedmontensis  will  be  used. 

Distribution  —  Massachusetts  and  Connecticut  southward  to  Ala¬ 
bama  and  Mexico  (Evans,  1930):  Temperate  element,  Appalachian  sub¬ 
element  (?),  Appalachian  unit  (?);  endemic. 

Subsection  Foliosae  (Bagl.  &  Carest.)  Vain. 

92.  Cladonia  robbinsii  Evans,  Trans.  Conn.  Acad.  Arts  Sci.  35: 
611.  1944. 

Material  seen  —  SUFFOLK  COUNTY:  Southold,  Latham  7550 
(+  7581),  January  20,  1934  (Latham);  Orient,  Latham  8467,  May  5, 
1939  (Latham);  Orient,  West  Long  Beach,  Latham  22304,  22309, 
December  7,  1944  (Latham);  Shinnecock  Hills,  Latham  24964,  May  8, 
1946  (Latham). 

This  species  bears  many  similarities  to  closely  related  C.  strepsilis, 
but  differs  from  the  latter  in  color  (dark  yellowish  green  as  opposed  to 
olive  green)  and  in  chemistry  (usnic  and  barbatic  acids  present  rather 
than  baeomycic  acid  and  strepsilin). 

Distribution  —  Connecticut  (Evans,  1944),  Tennessee,  Black  Hills; 
endemic. 

93.  Cladonia  strepsilis  (Ach.)  Vain.  Act.  Soc.  Faun.  FI.  Fenn.  10: 
403.  1894.  Baeomyces  strepsilis  Ach.  Meth.  Lich.  Suppl.  52.  1803. 

Material  seen  —  NASSAU  COUNTY:  Brodo  539  (16),  2527  (5), 
3350  (8),  3515  (10).  SUFFOLK  COUNTY:  23  specimens  collected  by 
Imshaug  and/or  Brodo;  17  specimens  collected  by  Latham  (Latham); 
Orient  Point,  Latham,  1927  (NY);  Shinnecock  Hills,  R.  H.  Torrey,  1933 
(NY);  Rt.  1  12  north  of  Coram,  R.  FI.  Torrey,  1936  (NY). 

No  other  Cladonia  on  Long  Island  has  strepsilin  and  the  accompany¬ 
ing  C  +  green  medullary  reaction. 

This  species  is  fairly  common  on  waste  soil  and  sandy  roadbanks; 
it  is  occasionally  found  on  mossy  soil  (figure  72). 

Distribution  —  Eastern  United  States  southward  to  Mexico  (Sand¬ 
stede,  1939):  Temperate  element,  East  Temperate  subelement;  Europe; 
Asia. 


184  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Subsection  Podostelides  (Wallr.)  Vain. 

Series  Helopodium  (Ach.)  Vain. 

94.  Cladonia  capitata  (Michx.)  Spreng.  Syst.  Veg.,  ed.  16,  4:  271. 
1827.  Helopodium  capitation  Michx.  FI.  Bor.  Am.  2:  329.  1803. 

Material  seen  - —  NASSAU  COUNTY:  Valley  Stream,  E.  A.  Warner, 
November  17,  1900  (BKL).  SUFFOLK  COUNTY:  Imshaug  25556 
(52);  Brodo  59-127  (54),  59-188  (54),  59-206  (68),  615  (39),  710A 
(65),  748  (53),  943  (59),  1359  (65),  1571  (65),  2167  (99), 

2737  (111),  2742  (111),  2485  ( 23 ) ,  3069  ( 128) ;  25  specimens  collected 
by  Latham  (Latham);  Orient,  Latham  190,  July  4,  1914  (FH);  Green- 
port,  Latham  5,  1914  (FH);  Coram,  R.  H.  Torrey,  1936  (NY). 

Fink  (1935)  listed  this  species  under  the  name  Cladonia  mitrula 
Tuck,  in  W.  Dari. 

Cladonia  capitata  is  most  commonly  found  on  tree  bases  in  well- 
lighted  oak  forests,  but  sometimes  is  found  on  sandy  soil. 

Distribution  —  Eastern  United  States  and  Cuba  (map:  Sandstede, 
1938):  Temperate  element.  East  Temperate  subelement;  Europe  (Poelt, 
1963 ) ;  Asia. 

95.  Cladonia  cariosa  (Ach.)  Spreng.  Syst.  Veg.  ed.  16,  4:  272. 
1827.  Lichen  cariosus  Ach.  Lich.  Suec.  Prodr.  198.  1798. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk  Point,  R.  H. 
Torrey,  1933  (NY). 

Distribution  —  Vermont,  Connecticut,  Tennessee,  Michigan,  On¬ 
tario,  Indiana,  Minnesota,  Black  Hills,  Arizona,  Washington,  Alaska, 
Saskatchewan,  Manitoba,  Baflin  Island:  Arctic-boreal  element,  circum- 
horeal. 

96.  Cladonia  subcariosa  Nyl.  Flora  59:  560.  1876. 

Material  seen  —  QUEENS  COUNTY:  Brodo  520  (3).  NASSAU 
COUNTY:  Brodo  3190  (6),  3348  (8).  SUFFOLK  COUNTY:  Imshaug 
25619  (116),  25622  (116),  25624  (116),  25625  (116),  25629  (116); 
Brodo  59-35  (53),  59-168  (82),  1797  (127),  2073  (38),  4227  (20); 
11  specimens  collected  by  Latham  (Latham);  Southold,  R.  H.  Torrey, 
1933  (NY);  Montauk  Point,  R.  H.  Torrey,  1933  (NY);  ?,  Latham  (18?), 
1914  (FH). 

One  can  consider  C.  subcariosa  the  central  element  of  a  group  of 
closely  related  taxa  called  the  Cladonia  subcariosa  group.  Members  of 
this  group  are  morphologically  almost  indistinguishable,  but  show  some 
differences  in  distribution  and  chemistry.  Of  this  group,  C.  subcariosa 
contains  norstictic  acid,  C.  clavulifera  contains  fumarprotocetraric  acid, 
C.  brevis  contains  psoromic  acid,  and  C.  polycarpia  contains  atranorin. 
Cladonia  polycarpia,  which  is  not  found  on  Long  Island,  is  considered 
synonymous  with  C.  clavulifera  by  Mattick  (1940).  In  this  paper,  the 
first  three  species  will  be  recognized  although  there  is  considerable  ques¬ 
tion  as  to  whether  they  are  distinct  (Mozingo,  1961).  In  view  of  the  fact 
that  these  species  differ  little  in  their  morphology,  and  their  chemical 
components  are  closely  “related”  (all  being  /3-orsellic  acid  depsides  or 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


185 


depsidones  with  a  substantial  history  of  chemical  shifting  between  closely 
related  taxa)  it  might  be  better  to  consider  them  in  an  appropriate  in¬ 
fraspecific  rank.  Pending  further  study  of  the  morphology,  chemistry,  and 
phytogeography  of  members  of  the  C.  subcariosa  complex,  the  various 
"microspecies”  will  be  recognized. 

Cladonia  subcariosa  is  found  in  dry,  sandy  or  grassy  fields. 

Distribution  —  Eastern  United  States  (map:  Sandstede,  1938): 
Temperate  element,  East  Temperate  subelement;  Europe;  Asia. 

97.  Cladonia  clavulifera  Vain,  in  Robb.  Rhodora  26:  145.  1924. 

Material  seen  —  NASSAU  COUNTY:  Brodo  2529  (5),  3498 

(4),  3504  (10).  SUFFOLK  COUNTY:  18  specimens  collected  by 
Imshaug  and/or  Brodo;  Southold,  Latham  7995  (+  8004),  February 
11,  1938  (Latham);  Orient,  Long  Beach,  Latham  22288,  22291, 
22299,  December  7,  1944  (Latham);  Napeague,  Latham  22983 

(—  22986),  February  20,  1941  (Latham);  Amagansett,  Latham  25997, 
March  11,  1947  (Latham);  Noyack,  Latham  26518,  March  9,  1947 
(Latham);  Bridgehampton,  Latham  27050,  September  14,  1947  (Lath¬ 
am);  North  Sea,  Latham  28152,  May  16,  1955  (Latham);  Orient  Point, 
Latham  2,  January  9,  1911  (NYS);  near  Orient,  Latham  13  (FH); 
(locality  unknown)  Latham,  1914  (FH);  Montauk  Point,  R.  H.  Torrey, 
1933  NY);  Southold,  R.  H.  Torrey,  1936  (NY);  Pike's  Beach,  West- 
hampton,  R.  H.  Torrey,  1936  (NY);  Selden,  S.  Cain  356,  347,  June  30, 
1936  (NY). 

This  species  is  usually  found  on  exposed,  sandy  ground. 

Distribution  — -  Maine,  Mass.,  Connecticut,  New  Jersey,  Maryland, 
Washington,  D.C.,  Virginia  (Sandstede,  1939);  Vermont,  Tennessee, 
Oklahoma:  Temperate  element.  East  Temperate  subelement(?) ;  Asia. 

98.  Cladonia  brevis  Sandst.  Ahhandl.  Naturv.  Ver.  Bremen.  25: 
192.  1922. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25666  (64); 
Brodo  1642  (69);  Southold,  Latham  7883,  February  1  1,  1938  (Latham); 
Riverhead,  Peck  (NYS);  Montauk  Point,  R.  H.  Torrey,  1933  (NY);  East 
of  Calverton,  R.  H.  Torrey  1936  (NY);  Airport  near  Westhampton, 
R.  H.  Torrey,  1936  (NY). 

Cladonia  brevis,  like  the  other  members  of  the  C.  subcariosa  group 
( p.  185)  is  found  on  dry  sandy  soil. 

Distribution  —  Maine,  Massachusetts,  Connecticut  (Sandstede, 
1938);  Vermont,  New  Jersey,  Tennessee,  Manitoba:  Temperate  element, 
East  Temperate  subelement  (?);  Europe. 

Subsection  Thallostelides  Vain. 

99.  Cladonia  verticillata  (Hoffm.)  Schaer.  Lich.  Helv.  Spic.  31. 
1823.  Cladonia  pyxidata  *  C.  verticillata  Hoffm.  Deutschl.  FI.  2:  122. 
1796. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  G.  B.  Brainerd, 
(1866?)  (BKL  031990);  New  Lots,  G.  B.  Brainerd,  (1866?)  (BKL). 
SUFFOLK  COUNTY:  Brodo  59-303  (53);  Greenport,  Latham  23430 


186  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

April  12,  1945  (Latham);  Northwest,  Third  station,  Latham  27447, 
April  27,  1948  (Latham);  Orient,  Latham  35337,  April  17,  1950  (Lath¬ 
am);  Sag  Harbor,  Latham,  September  15,  1941  (Latham);  ?  near  Orient, 
Latham  13  (FH). 

The  separation  of  this  species  from  closely  related  C.  calycantha  is 
often  very  difficult.  Such  characters  as  smooth  cup  margins  and  gradually 
expanding  cups  usually  attributed  to  C.  verticillata  are  not  always  evident. 
The  ecology  of  the  two  species,  however,  seems  to  be  different  with 
C.  verticillata  being  found  in  open,  sandy  or  grassy  fields,  especially  on 
neutral  soils,  and  C.  calycantha  being  found  mainly  in  boggy  or  acid  sand 
localities,  usually  under  pines  (figure  48).  The  geographical  distribution 
of  the  two  species  is  basically  different  as  well. 

Distribution  —  Nova  Scotia,  Vermont,  Massachusetts,  Connecticut, 
New  Jersey,  Tennessee,  Michigan,  Ontario,  Minnesota,  Black  Hills, 
Washington,  Alaska,  Saskatchewan,  Manitoba,  Canadian  East  Arctic; 
Arctic-boreal  element;  circumboreal. 

100.  Cladonia  calycantha  Nyl.  Syn.  Meth.  Lich.  192.  1858. 

Material  seen  —  SUFFOLK  COUNTY:  lmshaug  25856  (60); 

Brodo  59-23  (83 ),  59-25  (83 ),  59-306  (68),  59-307  (68),  1129  (78), 
2092  (83),  2287  (87),  2534  (49),  3396  (75),  3816  (66);  29  specimens 
collected  by  Latham  (Latham);  Napeague,  Latham  26024,  March  11, 
1947  (US:  Evans);  Springs,  Latham  26432,  April  17,  1947  (US:  Evans); 
Northwest,  Latham  26391,  April  17,  1947  (FH);  Northwest  Section  2, 
Latham  27480,  April  21,  1948  (US:  Evans);  Airport  near  Westhampton, 
R.  H.  Torrey,  1936  (NY);  Pike’s  Beach,  Westhampton,  R.  H.  Torrey, 
1936  (NY);  Sweezy  Pond,  2.3  miles  SW  of  Riverhead,  S.  Smith  11855, 
August  14,  1952  (NYS). 

The  relationship  between  this  species  and  C.  verticillata  has  been 
discussed  with  the  latter. 

Distribution  —  Newfoundland  to  Florida,  South  America,  Australia 
(Sandstede,  1938):  Tropical  element.  Coastal  Plain  subelement;  Europe 
(  Poelt,  1963 );  Asia. 

101.  Cladonia  mateocyatha  Robb.  Rhodora  27:  50.  1925. 

Material  seen  —  NASSAU  COUNTY:  Brodo  540  (  16).  SUFFOLK 

COUNTY:  ?,  Latham  34,  1914  (FH):  Montauk  Point,  R.  H.  Torrey ,  1933 
(NY);  between  Commack  and  Kings  Park,  R.  H.  Torrey,  1937  (NY). 

This  species,  very  rare  on  Long  Island,  shows  considerable  morpho¬ 
logical  variability.  Its  smooth  or  cracked,  completely  corticate  podetial 
surface,  together  with  its  irregularly  proliferating  cups,  giving  rise  to 
contorted  branches  from  both  central  and  marginal  areas,  distinguish 
C.  mateocyatha  from  other  nonsorediate  species. 

This  species  is  found  on  exposed  soil. 

Distribution  —  Massachusetts,  Connecticut,  Washington,  D.C.,  West 
Virginia,  New  Mexico  (Sandstede,  1939);  Vermont,  New  Jersey,  Tennes¬ 
see,  Michigan:  Temperate  element,  Appalachian  subelement,  Appalachian- 
Great  Lakes-Rocky  Mountain  unit  (?);  endemic. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  187 

102.  Cladonia  pyxidata  (L.)  Hoffm.  Deutschl.  FI.  2:  121.  1796. 
Lichen  pyxiclatiis  L.  Sp.  PI.  2:  1151.  1753. 

Material  seen  —  KINGS  COUNTY:  Gowanus,  G.  B.  Brainerd 
(  BKL)  031992).  SUFFOLK  COUNTY:  Imshaug  25628  (116),  25862 
(60);  Brodo  2838  (115);  10  specimens  collected  by  Latham  (Latham); 
Montauk  Point,  R.  H.  Torrey,  1933  (NY). 

The  species  is  usually  found  on  the  ground  in  dry,  sandy  localities. 

Distribution  ■ —  Vermont,  Massachusetts,  Connecticut,  New  Jersey, 
Tennessee,  Michigan,  Ontario,  Minnesota,  Wisconsin,  Black  Hills,  Ari¬ 
zona,  Idaho,  Washington,  Alaska,  British  Columbia,  Saskatchewan,  Mani¬ 
toba,  Can.  East  Arctic,  Baffin  Island:  Arctic-boreal  element;  circumboreal. 

103.  Cladonia  chlorophaea  (Florke  in  Somm.)  Spreng.  Syst.  Veg. 
ed.  16.  4:  273.  1827.  Cenomyce  chlorophaea  Florke  in  Somm.  Suppl.  FI. 
Lapp.  130.  1826. 

Material  seen  —  COUNTY  UNKNOWN:  Fresh  Pond,  Hulst,  1890 
(BKL  031986).  QUEENS  COUNTY:  Brodo  519  (3).  NASSAU 
COUNTY:  Brodo  537  (16),  541  (12),  560  (13),  1306  (15), 
3343  (8),  3507  (10);  Oyster  Bay,  L.  P.  le  ?,  September  1889  (NY); 
Valley  Stream,  Warner ,  November  17,  1900  (BKL).  SUFFOLK 
COUNTY:  109  specimens  collected  by  Imshaug  and/or  Brodo;  70  speci¬ 
mens  collected  by  Latham  (Latham);  (locality  unknown),  Latham,  1914 
(FH);  Orient,  Latham  185,  1914  (FH);  (locality  unknown),  Latham, 
1914  (FH);  Orient,  Latham  216,  May  23,  1914  (FH);  Shinnecock  Hills, 
R.  H.  Torrey,  1933  (NY);  Holtsville,  R.  H.  Torrey,  1936  (NY);  Pikes 
Beach,  Westhampton,  R.  H.  Torrey,  1936  (NY)  (PD  +  red);  Pikes 
Beach,  Westhampton,  R.  H.  Torrey,  1936  (NY)  (PD  —  );  Suffolk  County 
Airport  near  Westhampton,  R.  H.  Torrey,  1936  (NY);  Selden,  S.  Cain 
345,  June  30,  1936  (NY);  Wyandanch  Club  Game  Preserve  south,  R.  H. 
Torrey,  1937  (NY);  Horton’s  Beach,  Southold,  S.  Smith  11896,  11897, 
August  14,  1952  (NYS);  Wildwood  State  Park  near  Riverhead,  S.  Smith 
12744,  October  17,  1952. 

This  species,  one  of  the  most  abundant  on  Long  Island,  is  extremely 
variable  in  morpho'ogy  and  ecology.  Soredia  range  from  almost  farinose 
to  granular  and  even  appear  corticate  in  some  specimens;  cups  are  either 
simple,  goblet-shaped  structures  with  smooth  margins,  or  have  many 
often  large,  marginal  proliferations  bearing  large  brown  apothecia. 

Fumarprotocetraric  acid,  as  determined  by  a  PD  +  red  reaction  on 
the  podetia,  was  demonstrated  in  about  60  percent  of  the  specimens.  The 
presence  of  grayanic  acid  was  determined  by  the  microscopic  examina¬ 
tion  of  acetone  extracts  of  the  podetia,  with  supplementary  recrystalliza¬ 
tion  in  GAW  solution  if  necessary.  It  was  found  in  all  the  Long  Island 
specimens  except  one  ( Brodo  1050  [112])  in  which  cryptochlorophaeic 
acid  was  demonstrated.  In  addition,  9  specimens  of  C.  chlorophaea  from 
southern  New  Jersey  and  12  from  Cape  Cod  were  chemically  examined. 
All  of  these  also  contained  only  grayanic  acid  except  for  one  specimen 
( Brodo  4387  from  Cape  Cod)  which  showed  neither  grayanic  nor  crypto- 


188  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

chlorophaeic  nor  merochlorophaeic  acids. 

Fumarprotocetraric  acid  is  regarded  as  an  accessory  substance  in 
C.  chlorophaea  by  most  modern  workers  (see  Evans,  1944).  However, 
the  presence  or  absence  of  the  other  substances  mentioned  above  have 
been  used  by  Evans,  Asahina,  and  others  as  a  basis  for  recognizing  four 
species:  C.  grayi  Merr.  in  Sandst.  with  grayanic  acid;  C.  cryptochloro- 
phaea  Asah.  with  cryptochlorophaeic  acid;  C.  merochlorophaea  Asah. 
with  merochlorophaeic  acid;  C.  chlorophaea  sens.  str.  with  none  of  these 
chemicals.  Several  of  these  chemical  segregates  seem  to  have  some  geo¬ 
graphic  restrictions  (grayanic  acid  strain  is  eastern;  inactive  strain  is 
northern ) .  The  other  strains  are  rather  rare,  and  are  poorly  defined  geo¬ 
graphically.  Until  the  full  chemical  story  is  known,  at  least  in  North 
America,  it  seems  best  to  regard  these  segregates  as  chemical  strains 
within  C.  chlorophaea  sens,  lat.,  although  with  further  study  at  least  a 
few  may  prove  to  be  more  logically  considered  as  subspecies,  or  perhaps 
even  species. 

Cladonia  chlorophaea  is  found  on  soil,  lignum,  or  tree  bases. 

Distribution  —  Nova  Scotia,  Maine,  Vermont,  Massachusetts,  Con¬ 
necticut,  New  Jersey,  Tennessee,  Alabama,  Florida,  Michigan,  Ontario, 
Indiana,  Wisconsin,  Minnesota,  Oklahoma,  Black  Hills,  Arizona,  Wash¬ 
ington,  Alaska,  British  Columbia,  Manitoba,  Canadian  East  Arctic,  Baffin 
Island:  Arctic-boreal  element;  circumboreal. 

[Note:  Ahti’s  important  treatment  of  the  C.  chlorophaea  complex 
(Ahti,  1966)  appeared  after  this  paper  was  in  proof,  and  therefore  was 
not  considered  here.] 

104.  Cladonia  fimbriata  (L.)  Fr.  Lich.  Eur.  Ref.  222.  1831.  Lichen 
fimbriatus  L.  Sp.  PI.  1  152.  1753. 

Material  seen  —  SUFFOLK  COUNTY :  Riverhead,  Latham  7663B. 
May  1,  1937  (Latham);  Bridgehampton,  Latham  27043,  September  14, 
1947  (Latham);  North  Sea,  Latham  32353,  April  26,  1954  (Latham); 
Riverhead,  Latham  33318,  June  1,  1923  (Latham);  East  Marion,  Latham, 
May  3,  1914  (Latham). 

Cladonia  fimhriata  varies  from  a  very  narrow-cupped  condition  very 
similar  to  C.  coniocraea  to  a  broader  trumpet-shaped  condition  resembling 
C.  conista.  However,  its  podetia  always  show  a  distinct,  deep,  though 
often  very  narrow  cup  and  rarely  are  as  subulate  as  those  of  C.  conio¬ 
craea,  in  which  the  podetial  cups  are  flat  or  very  shallow.  Cladonia  fim¬ 
briata  also  has  much  smaller  podetial  and  basal  squamules  than  the  latter. 
In  addition,  the  podetia  of  C.  fimbriata  never  arise  from  the  center  of  a 
primary  squamule  as  do  the  podetia  of  C.  coniocraea.  Cladonia  conista, 
with  broad,  goblet-shaped  cups  has  its  soredia  confined  to  the  upper 
third  of  the  podetium  and  the  inner  surface  of  the  cup  and  contains 
substance  “H,”  whereas  C.  fimbriata  has  narrower  podetia  covered  with 
soredia  and  does  not  contain  substance  H. 

Cladonia  fimbriata  is  found  on  the  ground  and  on  tree  bases. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  189 

Distribution  —  Arctic-boreal  element  (Hale,  1954;  Thomson,  1953, 
1955):  circumboreal. 

105.  Cladonia  conista  (Ach.)  Robb,  in  Allen,  Rhodora  32:  92. 
1930.  Cenomyce  funbriata  (3-C.  conista  Ach.  Syn.  Meth.  Lich.  257.  1814. 

Material  seen  —  KINGS  COUNTY:  Gowanus,  G.  B.  Brainerd,  1866 
(BKL).  QUEENS  COUNTY:  Cypress  Hills,  Hulst,  1890  (BKL  031991). 
SUFFOLK  COUNTY:  Imshaug  25686  (72),  25750  (132);  Brodo  811 
(90B),  1523  (100B),  2725  (111),  3086  (128);  10  specimens  collected 
by  Latham  (Latham). 

The  presence  of  substance  H  in  C.  conista  easily  separates  it  from 
similar  species  which  lack  it,  such  as  C.  funbriata  (see  above)  and  C. 
chlorophaea.  In  addition,  the  latter  usually  has  distinctly  granular  soredia 
covering  the  entire  podetium. 

Cladonia  conista  grows  on  soil  or  tree  bases  (figure  69). 

Distribution  —  Vermont,  Massachusetts,  Connecticut,  New  Jersey, 
Tennessee,  Michigan,  Black  Hills:  Temperate  element,  North  Temperate 
subelement  (?);  Europe,  Asia. 

106.  Cladonia  conioeraea  (Florke)  Spreng.  era.  Sandst.  Syst.  Veg. 
ed.  16.  4:  272.  1827.  Sandstede,  Abh.  Naturw.  Ver.  Bremen  21:  373. 
1912.  Cenomyce  conioeraea  Florke,  Deutschl.  Lich.  138.  1821. 

Material  seen  —  QUEENS  COUNTY:  Brodo  526  (3).  NASSAU 
COUNTY:  17  specimens  collected  by  Brodo.  SUFFOLK  COUNTY: 
83  specimens  collected  by  Imshaug  and/or  Brodo;  47  specimens  collected 
by  Latham  (Latham);  Montauk  Point,  R.  H.  Torrey,  1933  (NY);  Coram, 
R.  H.  Torrey,  1936  (NY);  Greenport,  Latham  30938a,  May  30,  1952 
(NYS). 

This  species  is  one  of  the  most  common  and  variable  of  the  Cla- 
doniae.  Cladonia  conioeraea  is  usually  said  to  have  abruptly  tapering 
podetia  entirely  covered  with  farinose  soredia  except  for  a  narrow  basal 
zone.  Long  Island  specimens,  however,  show  every  gradation  from  this 
“typical”  form  to  a  condition  having  almost  entirely  corticate  podetia 
with  patches  of  farinose  or  granular  soredia  scattered  along  their  length. 
This  latter  form  has  generally  been  considered  under  the  name  Cladonia 
ochrochlora  Florke.  Evans  (1935*  discussed  the  difference  between  these 
two  species. 

In  general,  five  characters  are  fairly  constant  in  C.  conioeraea :  (1) 
the  podetia  are  sorediate,  (2)  the  podetia  are  usually  short,  stout,  and 
abruptly  tapering  to  a  sharp  point,  (3)  the  podetia  arise  from  the  center 
or  near  center  of  a  primary  squamule  (4)  the  primary  squamules  are 
broad,  and  (5)  the  podetial  cortex  anu  the  soredia  usually  have  a  yellow¬ 
ish  or  yellow-olive  cast  (not  due  to  usnic  acid). 

A  great  deal  of  variability  can  be  seen  in  (1)  the  extent  of  podetial 
cortex,  (2)  the  type  of  soredia  (although  the  granular  sorediate  condi¬ 
tion  is  very  rare),  (3)  the  presence  of  soredia  on  the  primary  squamules 
(from  abundant  to  essentially  absent),  (4)  the  presence  of  cupped 
podetia  (cf.  C.  funbriata,  p.  188),  (5)  the  lobing  of  the  primary  squam- 


190  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

ules  (entire  to  crenate),  and  (6)  the  degree  of  branching  ( podetia  are 
almost  always  simple,  but  rarely  have  one,  or  at  most,  two  simple 
branches).  Apothecia,  which  are  rare  in  this  species,  are  brown  and 
irregular,  occurring  at  the  edges  of  poorly  developed  cups  or  trays. 

Cladonia  coniocraea  is  usually  found  on  mossy  soil  or  tree  bases 
and  almost  always  in  shaded  situations.  Its  city-tolerance  is  discussed 
in  Brodo  ( 1966) . 

Distribution  —  Nova  Scotia,  Maine,  Vermont,  Massachusetts,  Con¬ 
necticut,  New  Jersey,  Tennessee,  Alabama,  Ontario,  Michigan,  Wisconsin, 
Minnesota,  Indiana,  Oklahoma,  Arizona,  Washington,  Alaska,  British 
Columbia,  Manitoba,  Canadian  East  Arctic:  Temperate  element  (?), 
North  Temperate  subelement;  Europe,  Asia. 

107.  Cladonia  nemoxyna  (Ach.)  Arn.  Lich.  exs.  no.  1495.  1890. 
Baeomyces  radiatus  /3.  B.  nemoxynus  Ach.  Meth.  Lich.  342.  1803. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2713  (111);  Orient, 
Long  Beach,  Latham  19,  April  15,  1914  (Latham);  Montauk  Point, 
R.  H.  Torrey,  1933  (NY);  Southold.  R.  H.  Torrey,  1936  (NY);  2.3  miles 
southwest  of  Riverhead,  S.  Smith  and  Ogden,  Smith  11853,  11854 , 
August  14,  1952  (NYS). 

Cladonia  nemoxyna  may  contain  fumarprotocetraric  acid;  it  always 
contains  homosekikaic  acid  (Evans,  1944).  All  the  Long  Island  speci¬ 
mens  were  PD  +  red  and  presumably  contained  fumarprotocetraric  acid. 
The  specimens  of  this  species  cited  by  Degelius  (1940)  from  Maine  also 
were  PD  +  red.  Specimens  from  Ontario  ( Ahti,  1964),  the  Great 
Lakes  region,  and  the  Black  Hills  are  PD—.  The  presence  of  fumar¬ 
protocetraric  acid,  therefore,  may  prove  to  have  geographic  correlation, 
as  do  many  other  chemical  populations  of  lichens  (Cladonia  uncialis, 
C.  chlorophaea,  Lecanora  caesiorubella,  etc.) 

Homosekikaic  acid  is  very  difficult  to  demonstrate,  apparently  be¬ 
cause  it  occurs  in  very  minute  concentrations.  The  directions  for  its 
recrystallization  from  GAoT  solution  as  given  by  Evans  (1943)  should 
be  followed  carefully. 

This  species  was  found  on  eroding  soil  associated  with  C.  conista, 
C.  farinacea,  and  C.  cristatella. 

Distribution  —  Vermont,  Massachusetts,  Connecticut,  New  Jersey, 
Tennessee,  Ontario,  Michigan,  Black  Hills,  Washington:  Temperate  ele¬ 
ment,  North  Temperate  subelement;  Europe;  Asia. 

108.  Cladonia  cylindrica  (Evans)  Evans,  Rhodora  52:  116.  1950. 
Cladonia  borhonica  f.  cylindrica  Evans,  Trans.  Conn.  Acad.  Arts  Sci. 
30:  482.  1930. 

Material  seen  —  NASSAU  COUNTY:  Brodo  1309  (15),  3500 
(10).  SUFFOLK  COUNTY:  Brodo  711 A  (65),  955  (35-36),  1012 
(27),  1262  (29),  1700  (133),  2226  (28),  3909  (112);  Greenport,  Gull 
Pond,  Latham  29595,  January  20,  1951,  dry  soil  in  woods  (Latham). 

This  lichen  bears  many  similarities  with  C.  coniocraea.  Both  have 
more  or  less  short,  sorediate,  usually  sterile  podetia,  and  both  are  PD  + 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


191 


red.  However,  C.  cylindrica  has  a  clear  gradation  of  coarse  granules  at 
the  podetial  base  to  farinose  soredia  at  its  tip  and  contains  grayanic  acid, 
whereas  C.  coniocraea  is  entirely  covered  with  farinose  soredia  and  lacks 
grayanic  acid. 

Cladonia  cylindrica  is  found  on  tree  bases,  usually  in  shaded  woods. 

Distribution  —  Vermont,  Massachusetts,  Connecticut,  New  Jersey, 
Michigan;  West  Virginia  (Sandstede,  1939):  Tropical  element,  Tem- 
perate-Appalachian  subelement;  Asia;  circumtropic  (Sandstede,  1939). 

109.  C'adonia  pityrea  (Florke)  Fr.  Nov.  Sched.  Crit.  21.  1826. 
Capitularia  pityrea  Florke,  Ges.  Naturf.  Fr.  Berlin  Mag.  2:  135.  1808. 

Material  seen  —  QUEENS  COUNTY:  Brodo  523  (3).  SUFFOLK 
COUNTY:  Brodo  585  (92),  2747  (111),  3266  (119),  3323  (129); 
East  Marion,  Latham  5,  May  3,  1914  (Latham);  Greenport,  Latham 
23428,  April  12,  1945  (Latham);  Riverhead,  Latham  36811 A,  May  1, 
1960  (Latham);  Riverhead,  Latham,  May  25,  1960  (Latham);  Green- 
port,  Latham  7212  (?)  (MICH);  (locality  unknown),  Latham,  1914 
(FH). 

Almost  all  of  the  Long  Island  specimens  of  C.  pityrea  are  identical 
with  Connecticut  material  identified  by  Evans  (in  herb.  FH)  as  C.  pityrea 
var.  zwackii  Vain.,  and  either  form  subacuta  Vain,  or  form  squamulifera 
Vain.  The  podetia  were  contorted  and  covered  with  coarse  granules  or 
granular  soredia.  In  the  type  of  form  squamulifera  ( Thaxter  35,  Trinidad, 
1912-13,  [FH]),  the  podetia  were  densely  squamulose  and  granular 
sorediate,  not  very  contorted,  and  were  not  pellucid  and  dark  in  the 
decorticate  areas.  The  podetial  squamules  were  finely  lobed  and  almost 
nonsorediate.  In  other  words,  the  type  of  f.  squamulifera  does  not  seem 
to  agree  with  Evans’  identifications,  and  /.  squamulifera  sensu  Evans  is 
probably  a  kind  of  /.  subacuta  (especially  since  collections  containing  both 
forms  in  the  same  packet  were  common).  The  value  of  these  infra¬ 
specific  taxa  is  doubtful. 

Cladonia  pityrea  has  been  collected  on  various  substrates,  including 
dry  ground,  tree  bases,  rocks,  and  wood. 

Distribution  —  Vermont,  Connecticut,  New  Jersey,  Tennessee,  Flor¬ 
ida,  Michigan;  South  America,  West  Indies,  East  Indies  (Sandstede, 
1939):  Tropical  element,  Appalachian-Temperate  subelement;  Europe; 
Asia. 

110.  Cladonia  simulata  Robb.  Rhodora  31:  105.  1929. 

Material  seen  —  SUFFOLK  COUNTY:  Northwest,  Latham  27200 , 

April  27,  1947  (US:  Evans). 

This  species  had  been  placed  in  the  subsection  Ochroleucae  by 
Sandstede  (1938)  and  Mattick  (1940)  on  the  basis  of  its  resemblance  to 
C.  piedmontensis.  Evans  (1952),  following  Robbins,  pointed  out  that  the 
similarity  of  the  species  to  C.  piedmontensis  is  entirely  superficial,  and  its 
chemistry  (lack  of  usnic  and  presence  of  fumarprotocetraric  acid)  places 
it  in  the  subsection  Thallostelides,  close  to  C.  pityrea. 

The  specimen  was  growing  on  dry  sand. 


192  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Distribution  —  Massachusetts,  North  Carolina,  Georgia,  Florida 
(Evans,  1952):  Temperate  element,  Coastal  Plain  subelement;  endemic. 

Section  PERVIAE  (Fr. )  Matt. 

Subsection  Chasmariae  (Ach.)  Florke 
Series  Megaphyllae  Vain. 

111.  Cladonia  apodocarpa  Robb.  Rhodora  27:  211.  1925. 

Material  seen  —  SUFFOLK  COUNTY:  Riverhead,  Latham  24794 r 

April  5,  1946  (US:  Evans). 

This  species  is  the  only  nonpodetiate  Cladonia  on  Long  Island, 
containing  both  atranorin  and  fumarprotocetraric  acid.  Latham’s  speci¬ 
men  was  from  a  dry  woods. 

Distribution  —  Northeastern  states  (Sandstede,  1939);  Tennessee, 
Alabama:  Temperate  element,  East  Temperate  subelement;  endemic. 

Series  Microphyllae  Vain. 

112.  Cladonia  caespiticia  (Pers.)  Florke,  Clad.  Comm.  8.  1828. 
Baeomyces  caespiticus  Pers.  Ann.  d.  Rot.  7:  155.  1794. 

Material  seen  —  QUEENS  COUNTY:  Brodo  521  (3).  NASSAU 
COUNTY:  Brodo  1310  (15),  2528  (5).  SUFFOLK  COUNTY:  Imshaug 
25693  (72) ;  Brodo  2426  (20),  2755  (107),  3023  (50),  3864  (57), 
3912  (36);  Orient,  Latham  789,  March  3,  1914  (Latham);  Montauk, 
Latham  31892,  May  16,  1951  (Latham);  Mattituck,  Latham  33140, 
June  7,  1955  (Latham);  East  Marion,  Latham,  May  3,  1914  (Latham); 
Orient,  Latham  199,  May  15,  1914  (FH);  (locality  unknown),  Latham, 
1914  (FH);  Barling  Hollow  (=  Baiting  Hollow?),  R.  H.  Torrey,  1934 
(NY);  Wyandanch  Club  Game  Preserve,  south  of  Smithtown,  R.  H. 
Torrey,  1937  (NY). 

This  species  is  found  on  bare  or  mossy  ground,  often  on  charred 
ground,  and  less  frequently  on  tree  bases.  It  is  almost  always  in  shaded  or 
partialiy  shaded  localities,  particularly  in  the  black  and  red  oak  forests 
(figure  55). 

Distribution  —  Eastern  United  States  (Sandstede,  1938) :  Temperate 
element,  East  Temperate  subelement;  Europe;  Asia. 

113.  Cladonia  parasitica  (Hoffm.)  Hoffm.  Deutschl.  FI.  2:  127, 
1795.  Lichen  parasiticus  Hoffm.  Enum.  Lich.  39,  tab.  8,  fig.  5.  1784. 

Material  seen — NASSAU  COUNTYs  Brodo  1505  (14),  1507  (14). 
SUFFOLK  COUNTY:  Brodo  927  (59),  127 3 A  (21),  1293  (19),  1296 
(19),  2100  (78),  2153  (102),  2456  (22),  2643  (71),  2975  (43),  2978 
(43),  3041  (50),  3155  (65),  3878  (62)  3906  (112);  12  specimens 

collected  by  Latham  (Latham);  Shinnecock  Hills,  R.  H.  Torrey,  1933 
(NY). 

Cladonia  parasitica  is  usually  described  as  having  granular  soredia. 
On  many  specimens  which  I  have  seen,  however,  the  so-called  soredia 
appear  to  be  corticate  and,  therefore,  are  actually  granules. 

The  species  has  been  called  C.  delicata  (Ehrh.)  Florke  by  most 
authors. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


193 


This  species  is  almost  entirely  confined  to  decaying  logs  and  wood 
of  various  origins,  but  especially  coniferous  trees.  It  is  usually  found 
in  shaded  areas. 

Distribution  —  Throughout  eastern  United  States  (Evans,  193C): 
Temperate  element,  East  Temperate  subelement;  Europe;  Asia. 

114.  Cladonia  santensis  Tuck.  Amer.  Sci.  Arts  II.  25:  427.  1858. 
Material  seen  —  SUFFOLK  CO’  NTY:  hnshaug  25820  (86); 

Riverhead,  Sweezy  Pond,  Latham  32t-*U,  April  IS,  1955,  (Latham); 
North  Sea,  Latham  36939,  May  20,  1954  (Latham);  Riverhead,  Latham, 
May  1,  1960  (Latham). 

This  species  can  be  recognized  even  in  the  field  by  its  very  pale, 
almost  white  color,  and  its  contorted,  bent,  minutely  squamulose  podetia. 
Sterile  material  of  C.  santensis  sometimes  closely  resembles  sterile  C.  par¬ 
asitica,  which  also  contains  thamnolic  acid.  The  latter,  however,  has 
finely  divided,  “lacy,”  granulate  primary  squamules,  whereas  in  C.  san¬ 
tensis  the  squamules  are  thicker  and  not  granular  or  sorediate. 

Long  Island  is  the  northern  limit  of  this  species  (figure  37).  It  was 
found  to  be  abundant  in  cedar  bogs  in  southern  New  Jersey. 

Distribution  —  New  Jersey  to  Florida  (Evans,  1952):  Temperate 
element,  Coastal  Plain  subelement;  endemic. 

115.  Cladonia  squamosa  (Scop.)  Hoffm.  Deutschl.  FI.  2:  125.  1796. 
Lichen  squamosus  Scop.  FI.  Carn.  ed.  2.  2:  368.  1772. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2164  (102),  2358 
(42),  2569  (73),  2998  (17),  3001  (17),  3003  (17),  3035  (50);  10 
specimens  collected  by  Latham  (Latham). 

This  species,  which  is  uncommon  on  Long  Island,  is  usually  found 
on  mossy  ground,  rocks,  or  woods  in  partially  shaded  black  or  red  oak 
forests  (figure  54).  Its  relationship  to  C.  atlantica  is  discussed  under  the 
latter  species. 

Distribution  —  Nova  Scotia,  Maine,  Vermont,  Massachusetts,  Con¬ 
necticut,  New  Jersey,  Tennessee,  Alabama,  Ontario,  Michigan,  Minne¬ 
sota,  Indiana.  Washington,  coastal  Alaska,  Baffin  Island;  “arctic  to  south¬ 
ern  temperate”  (Ahti,  1964):  Arctic-boreal  element;  circumboreal. 

116.  Cladonia  atlantica  Evans,  Trans.  Conn.  Acad.  Arts  Sci.  35: 
573.  1944. 

Material  seen  —  NASSAU  COUNTY:  Brodo  542  (12),  1312  (15), 
3502  (10).  SUFFOLK  COUNTY:  59  specimens  collected  by  Imshaug 
and/or  Brodo;  66  specimens  collected  by  Latham  (Latham);  West  Suffolk 
Co.  Airport  near  Westhampton,  R.  H.  Torrey  (NY);  Baiting  Hollow, 

R.  H.  Torrey,  1934  (NY);  Holtsville,  R.  H.  Torrey,  1936  (NY);  Smith- 
town,  R.  H.  Torrey,  1936  (NY);  2.3  miles  SW  of  Riverhead,  S.  Smith 
11852,  11856,  11857,  August  14,  1952  (NYS);  Horton’s  Beach,  Southold, 

S.  Smith  11895,  August  14,  1952  (NYS);  1.3  miles  W  of  Middle  Island, 
S.  Smith  17717,  March  12,  1955  (NYS). 

This  species,  which  is  very  common  throughout  the  sandy  parts  of 
the  island,  is  very  variable  in  its  morphology.  Podetia  devoid  of  squam- 


194  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

ules  commonly  are  found,  as  well  as  podetia  entirely  covered  with  small 
or  large  squamules.  Apothecia  seem  to  be  more  common  on  the  squamu- 
lose  forms. 

The  main  difference  between  C.  atlantica  and  C.  squamosa  (from 
which  it  was  segregated  by  Evans)  is  in  the  production  of  baeomycic 
acid  in  the  former.  Evans  (1944)  discussed  their  differences  and  simi¬ 
larities  in  detail.  The  two  species  also  differ  in  ecology  and  distribution. 
Cladonia  atlantica  grows  on  acid  sand  and  lignum  and  is  more  or  less 
photophilous.  Cladonia  squamosa  is  a  species  of  partially  shaded,  mossy, 
rich  soil  habitats. 

Distribution  —  Temperate  element.  Coastal  Plain  subelement  (see 
Hale,  1961a);  endemic. 

117.  Cladonia  beaumontii  (Tuck.)  Vain.  Acta  Soc.  Faun.  FI.  Fenn. 
10:  455.  1894.  Cladonia  santensis  b.  beaumontii  Tuck.  Syn.  N.  Amer. 
Lich.  1:  245.  1882. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2249  (87);  21  speci¬ 
mens  collected  by  Latham  (Latham). 

This  species  is  very  closely  related  to  C.  atlantica  which,  however, 
always  shows  more  or  less  distinct  cups.  In  addition,  C.  beaumontii  is. 
usually  more  decorticate  than  C.  atlantica. 

Cladonia  beaumontii  is  a  lignum-inhabiting  bog  species  (figure  35). 
Distribution  —  Massachusetts,  Connecticut,  New  York,  North  Caro¬ 
lina,  Florida  (Evans,  1950):  Temperate  element.  Coastal  Plain  subele¬ 
ment;  endemic. 

118.  Cladonia  floridana  Vain,  in  Sandst.  Clad.  Exsic.  1196.  1922. 
Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25638  (64), 

25665  (64),  25827  (86);  Brodo  652  (79),  654  (79),  1151  (70),  1933 
(85),  1948  (85),  3393  (75),  3404  (75);  23  specimens  collected  by 
Latham  (Latham);  Suffolk  Co.  Airport  near  Westhampton,  R.  H.  Torrey, 

1936  (NY);  Rock  Hill  (near)  S.  of  Calverton,  Latham  7822,  June  28, 

1937  (NYS). 

Cladonia  floridana  is  found  on  exposed  or  partially  shaded  sand 
or,  rarely,  on  wood  (figure  19).  Although  it  is  almost  entirely  limited 
to  the  coastal  plain,  R.  H.  Torrey  (in  Smiley,  1940)  reported  its  occur¬ 
rence  in  Ellenville,  N.  Y.  (Ulster  County)  in  the  Shawangunk  Mountains 
at  an  elevation  of  2,200  feet.  This  unlikely  distribution  is  repeated  by  the 
heath  Corema  conradii,  which  is  typically  a  coastal  plain  species  but  is 
also  found  at  1500  feet  on  Gertrude’s  Nose,  also  in  the  Shawangunk 
mountains. 

Distribution  - — -  Cape  Cod  to  Florida  (Evans,  1952):  Temperate 
element,  Coastal  Plain  subelement;  endemic. 

119.  Cladonia  multiformis  Merr.  Bryologist  12:  1.  1909. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2703  (111);  (local¬ 
ity  unknown),  Latham  28,  1914  (FH). 

This  very  rare  species  was  found  on  dry  soil. 

Distribution  —  Nova  Scotia,  Vermont,  Connecticut,  Ontario,  Michi- 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  195 

gan.  Black  Hills,  Washington,  Saskatchewan,  Manitoba:  Temperate  ele¬ 
ment,  North  Temperate  subelement;  Africa  (des  Abbayes,  1938). 

120.  Cladonia  scabriuscula  (Del.  in  Duby)  Nyl.  Compt.  Rendu  83: 
88.  1876.  Cenomyce  scabriuscula  Del.  in  Duby,  Bot.  Gall.  632.  1830. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  G.  B.  Brainerd , 
1866  ( BKL) ;  New  Lots,  G.  B.  Brainerd,  1866  (BKL  031988).  SUF¬ 
FOLK  COUNTY:  Imshaug  25736  (132);  Brodo  59-276  (54),  1807 
(125),  2622  (84);  near  Orient,  Latham  2,  May  1914  (FH). 

This  species  and  its  relationship  to  C.  farinacea  is  discussed  in  detail 
with  the  latter.  Cladonia  scabriuscula  sens.  str.  is  very  rare  on  Long 
Island.  It  is  found  on  mossy  ground  and  on  tree  bases,  usually  in  the  shade. 

Distribution  —  Nova  Scotia,  Vermont,  Massachusetts,  Connecticut, 
Michigan,  Ontario,  Black  Hills,  coastal  Alaska;  arctic  to  southern  tem¬ 
perate,  with  oceanic  tendencies  (Ahti,  1964):  Arctic-boreal  element  (?); 
Europe;  Asia. 

121.  Cladonia  farinacea  (Vain.)  Evans,  Rhodora  52:  95.  1950. 
C.  furcata  y  scabriuscula  f.  farinacea  Vain.  Acta  Soc.  Faun.  FI.  Fenn. 
4:  339.  1887. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  G.  B.  Brainerd, 
(with  Cladonia  bacillaris )  (BKL  031984).  SUFFOLK  COUNTY:  Brodo 
59-276  (54),  59-296  (54),  1833  (125),  2712  (111),  3412  (134),  3183 
(72);  19  specimens  collected  by  Latham  (Latham). 

After  looking  at  many  specimens  of  both  C.  scabriuscula  and  C. 
farinacea  from  several  parts  of  the  country,  I  am  not  at  all  convinced 
that  the  two  are  actually  different  species. 

In  C.  scabriuscula,  the  podetia  are  typically  tall,  branched,  covered 
with  small  or  large  squamules  (often  sorediate)  and  become  granular 
sorediate  towards  their  tips.  The  squamules  are  often  very  inconspicuous 
on  the  upper  half  of  the  podetium,  and  the  granular  soredia  are  often 
abundant  over  the  greater  part  of  the  podetium. 

The  Long  Island  material  of  C.  scabriuscula  usually  is  short  (less 
than  20  mm  tall),  squamulose,  and  irregularly  sorediate  with  clumps  of 
granular  soredia. 

Cladonia  farinacea  typically  has  tall  podetia  which  are  infrequently 
branched,  farinose  sorediate  for  most  of  their  length,  and  almost  de¬ 
void  of  podetial  squamules. 

Long  Island  C.  farinacea,  however,  is  rather  short,  often  has  granu¬ 
lar  soredia,  and  occasionally  even  shows  some  podetial  squamules.  Evans 
identified  all  of  Latham’s  C.  scabriuscula  sens  lat.  as  C.  farinacea,  in¬ 
cluding  a  specimen  which  I  am  calling  C.  scabriuscula  sens.  str.  (Latham 
7522).  In  other  words,  Evans’  concept  of  C.  farinacea  was  apparently 
very  broad  and  allowed  for  considerable  variation  in  the  principal 
separating  characters. 

Cladonia  farinacea  is  usually  found  in  dry,  exposed,  grassy  fields  or 
on  eroded  ground.  Cladonia  scabriuscula  is  usually  on  richer  soil  in  more 
shaded  localities. 


196  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  distribution  of  the  two  species  seem  to  be  fairly  distinct  in 
most  areas. 

Distribution  —  Widely  distributed  in  North  America;  in  eastern 
part,  south  to  North  Carolina  and  west  to  Wisconsin  (Evans,  1950); 
Punta  Arenas,  southern  tip  of  Chile  (type  locality),  Port  Famine,  Straits 
of  Magellan  (Evans,  1950);  Asia. 

122.  Cladonia  furcata  (Huds.)  Schrad.  Spic.  FI.  Germ.  107.  1794. 
Lichen  furcatus  Huds.  FI.  Angl.  458.  1762. 

Material  seen  —  QUEENS  COUNTY:  Fresh  Pond,  Hulst,  1890, 
(BKL  031989).  SUFFOLK  COUNTY:  Imshaug  2561 31*  (116),  25618 
(116);  Brodo  59-174  (100B),  1166  (70),  1814  (125),  2046  (45), 
2613  (84),  2741  (111),  2746  (111),  3092  (126),  3185  (69),  3250 
(119),  3365  (94),  3370li  (94),  3890  (112);  52  specimens  collected 
by  Latham,  including  787014  (Shinnecock  Hills)  (Latham);  Northport, 
Grout,  December  1900  (BKL  031987);  ?,  Latham,  May  1914  (FH); 
Southold,  Latham  188,  October  4,  1914  (FH). 

Cladonia  furcata  shows  considerable  morphological  variation  with 
various  ecological  situations.  It  is  usually  pale  green  and  more  or  less 
squamulose  in  the  shade  on  mossy  banks,  and  is  slender,  distinctly 
browned,  and  essentially  devoid  of  squamules  in  fully  exposed  localities 

(p.  110). 

A  small  percentage  of  the  northeastern  material  of  C.  furcata  shows 
the  presence  of  atranorin,  including  several  specimens  from  Long  Island 
(see  above),  one  from  Nantucket  ( Brodo  4165)  and  one  from  Cape  Cod 
( Brodo  4330).  These  specimens  which  appear  like  C.  furcata  but  differ 
chemically  were  called  C.  subrangiformis  Sandst.  by  Evans  (1954). 
Ahti  (1962)  examined  the  type  of  the  latter  and  stated  that  it  seems  to 
be  distinct  from  C.  furcata.  He  believes  the  North  American  material 
with  atranorin,  however,  is  merely  a  chemical  race  of  C.  furcata. 

Cladonia  furcata  is  found  on  exposed  or  partially  shaded,  sandy  or 
grassy  ground  and,  rarely,  is  also  found  on  wood  or  mossy  boulders. 

Distribution  —  From  arctic  regions  southward  into  Mexico  (Evans, 
1930),  but  not  reported  by  Hale  (1954),  Thomson  (1953,  1955),  or 
Ahti  (1964):  Arctic-boreal  element  (?);  circumboreal. 

123.  Cladonia  carassensis  Vain.  Acta  Soc.  Faun.  FI.  Fenn.  4:  313. 

1887. 

Material  seen  —  SUFFOLK  COUNTY :  Three  Mile  Harbor,  Latham 
26432,  April  17,  1947  (Latham);  Riverhead,  Latham  30565,  April  3, 
1952  (Latham). 

The  Latham  specimens  were  found  on  rotten  wood  and  sandy  soil 
in  bogs  and  swamps.  Evans  (1950)  discusses  the  species  in  detail. 

Distribution  —  Massachusetts,  Connecticut,  Oregon,  Haiti,  Brazil, 
New  Zealand  (Evans,  1950):  Tropical  element,  Oceanic  subelement; 
eastern  Europe  (Evans,  1950);  Asia  (Asahina,  1950,  sub  C.  japonica 
Vain.) . 


“Contains  atranorin,  as  demonstrated  with  GAoT. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


197 


Subsection  Unciales  (Del.)  Vain. 

124.  Cladonia  boryi  Tuck.  Proc.  Amer.  Acad.  Arts  Sci.  1 :246.  1847. 
(Syn.  Lich.  New  Eng.  54.  1848.) 

Material  seen  —  NASSAU  COUNTY:  Meadowbrook  Valley, 
Hempstead  Plains,  Harper,  March  27,  1918  (NY).  SUFFOLK  COUN¬ 
TY:  28  specimens  collected  by  Imshaug  and/or  Brodo;  40  specimens 
collected  by  Latham  (Latham);  16  specimens  collected  by  R.  H. 
Torrey,  1933-1937  (NY);  Orient  Point,  (collector  unknown),  September 
1870  (FH);  Southampton,  Chite,  September  3-7,  1898  (NY);  Wading 
River,  Peck,  September  (NYS);  Wading  River,  Peck  (NYS);  Orient, 
Young  (BKL);  Orient  Point,  Latham,  December  8,  1909  (NYS);  Orient, 
Latham  1,  1913  (FH);  (locality  unknown),  Latham  30,  1914  (FH); 
Three  Mile  Harbor,  Latham  26412,  April  17,  1947  (FH);  Tiana  Beach, 
S.  Smith  28842,  August  4,  1959  (NYS);  3  miles  south  of  Montauk 
Point,  Gillis  4928,  September  7,  1961  (MSC). 

The  external  morphology  of  this  species  is  very  variable  and  one 
should  mainly  rely  on  the  internal  anatomy  described  in  the  key. 

Cladonia  boryi  is  strictly  an  exposed  sand  plain  and  sand  dune 
species  (figure  74). 

Distribution  - —  Nova  Scotia,  Massachusetts,  Connecticut,  New  Jer¬ 
sey:  Temperate  element,  Coastal  Plain  subelement  (?);  Asia  and  Brazil 
(Vainio,  1887). 

125.  Cladonia  caroliniana  Tuck.  Amer.  J.  Sci.  Arts  II.  25:  427. 
1858. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3346  (8),  3349  (8); 
Plain  Edge,  S.  Cain  373,  August  3,  1936,  A ndropogonetum  Hempsteadii 
(NY).  SUFFOLK  COUNTY:  25  specimens  collected  by  Imshaug  and/or 
Brodo;  38  specimens  collected  by  Latham  (Latham);  Coram,  N.  Taylor 
1,  June  15,  1922  (NY);  Selden,  S.  Cain  358,  June  30,  1936  (NY); 
Coram,  R.  H.  Torrey  (2  specimens),  1936  (NY);  Reeves  Bay  near 
Flanders,  R.  H.  Torrey,  1937  (NY);  Rt.  112  north  of  Coram,  R,  H. 
Torrey  (2  specimens),  1936  (NY);  Barling  Hollow  (Baiting  Hollow?), 
R.  H.  Torrey,  1937  (NY);  Pikes  Beach,  Westhampton,  R.  H.  Torrey, 
1936  (NY);  1.3  miles  W  of  Middle  Island,  S.  Smith  17716,  March  12, 
1955  (NYS);  Tiana  Beach,  S.  Smith  28443,  August  4,  1959  (NYS). 

Cladonia  caroliniana,  like  C.  uncialis,  is  found  on  sandy  or  mossy 
soil  in  exposed  or  partially  shaded  localities  (figure  76). 

Distribution  —  Throughout  eastern  United  States  (Evans,  1952): 
Temperate  element,  East  Temperate  subelement;  endemic. 

126.  Cladonia  uncialis  (L.)  G.  Web.  in  Wigg.  Primit.  FI.  Holsat. 
90.  1780.  Lichen  uncialis  L.  Sp.  PI.  1153.  1753. 

Material  seen  —  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 
(1866?)  (BKL).  NASSAU  COUNTY:  Meadow  Brook  Valley,  Hemp¬ 
stead  Plains,  Harper,  March  27,  1918  (NY).  SUFFOLK  COUNTY: 
36  specimens  collected  by  Imshaug  and/or  Brodo;  66  specimens  col¬ 
lected  by  Latham  (Latham);  Orient,  Young  (BKL);  Shinnecock  Hills, 


198 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


R.  H.  Toney,  1933  (NY);  Montauk  Point.  R.  H.  Toney,  1933  (NY); 
Airport  near  Westhampton.  R.  H.  Toney,  1936  (NY);  Coram.  R.  H. 
Torrey,  1936  (NY);  Selden,  5.  Cain  351,  June  30,  1936  (NY). 

Cladonia  uncialis  shows  several  growth  forms,  apparently  in  response 
to  different  ecological  situations.  In  exposed  areas  on  bare  sand,  the 
podetia  are  slender  and  crowded,  forming  tight,  flattened  cushions;  in 
shaded  localities  on  mossy  soil  or  in  protected  spots  where  moisture  is 
usually  abundant,  the  podetia  become  broad,  tall,  and  erect  without 
forming  distinct  cushions.  The  smooth,  somewhat  pruinose  podetial  inner 
lining,  however,  is  constant  for  the  species. 

The  chemistry  of  the  species  is  somewhat  variable,  with  squamatic 
acid  occurring  in  some  geographic  areas  and  not  in  others  (Evans,  1944). 
On  Long  Island,  all  specimens  have  a  medullary  white  UV  fluorescence 
and  all  those  extracted  with  acetone  and  tested  with  GE  solution  showed 
the  presence  of  squamatic  acid.  A  study  of  the  material  of  C.  uncialis  in 
the  Michigan  State  University  herbarium  revealed  that  the  squamatic 
acid  strain  is  found  in  the  Appalachian  Mountain  range  and  along  the 
northeast  coast  as  far  north  as  New  Brunswick  as  well  as  in  boreal  and 
arctic  Canada  and  Alaska.  The  squamatic  negative  strain  seems  to  be 
confined  to  the  Great  Lakes  region  and  northern  New  England.  In  Europe 
the  squamatic  strain  is  found  in  central  portions  of  the  continent  and 
the  inactive  strain  is  mainly  found  in  Scandinavia  and  Russia  (Evans, 
1944). 

Distribution  —  Arctic  regions  south  to  Alabama  (Evans,  1930): 
Arctic-boreal  element;  circumboreal. 

Subgenus  CLAD1NA  (Nyl.)  Leight.  em.  Vain. 

Section  BICORNUTAE  Abb. 

127.  Cladonia  evansii  Abb.  Lond.  J.  Bot.  76:  351.  1938. 

Material  seen  —  SUFFOLK  COUNTY:  Shinnecock,  Latham  33156, 
April  30,  1926.  (US:  Evans,  Latham). 

Latham’s  specimen  was  first  identified  as  Cladonia  impexa  f.  conden- 
sata  (Florke)  Sandst.  by  Evans,  who  noted  the  presence  of  usnic  acid, 
perlatolic  acid,  and  atranorin.  Ahti  and  Thomson  later  studied  the  same 
specimen  and  called  it  C.  evansii  (Ahti,  1961).  When  I  first  came  upon 
a  duplicate  of  the  specimen  in  the  Latham  herbarium.  I  referred  it  to 
C.  terrae-novae  Ahti,  having  demonstrated  atranorin  and  what  appeared 
to  be  usnic  acid  in  GAoT  solution.  However,  after  seeing  the  Evans 
herbarium  material  which  was  much  better  developed,  and  after  exam¬ 
ining  many  specimens  of  both  C.  terrae-novae,  which  I  collected  on 
Cape  Cod  and  Nantucket  Island,  and  C.  evansii  from  the  Michigan  State 
University  herbarium,  1  also  came  to  the  conclusion  that  the  Latham 
specimen  must  indeed  be  C.  evansii  with  Long  Island  thus  representing 
its  northernmost  locality.  The  differences  between  C.  evansii  and  C. 
terrae-novae  are  discussed  on  p.  201. 

I  atham’s  specimen  of  C.  evansii  was  found  on  dry  sandy  soil  on  an 
open  hill. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


199 


Distribution  —  Southeastern  United  States  and  the  West  Indies 
(map:  Ahti,  1961):  Temperate  element,  Coastal  Plain  subelement; 
endemic. 

Section  ALPESTRES  Abb. 

128.  Cladonia  alpestris  (L.)  Rabenh.,  Clad.  Europ.  11.  1860. 

Lichen  rangiferinus  (y)  alpestris  L.  Sp.  PI.  1153.  1753. 

Material  seen  —  KINGS  COUNTY:  Forest  Park,  Hulst,  November 
30,  1890  ( BKL  031993).  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brain- 
erd,  (1866?)  (BKL).  SUFFOLK  COUNTY:  Brodo  653  (79),  3887 
(120);  12  specimens  collected  by  Latham  (Latham). 

Cladonia  alpestris  at  one  time  was  probably  fairly  abundant  through¬ 
out  the  eastern  part  of  Long  Island  (Latham.  1949),  even  having 
occurred  in  the  New  York  City  area  at  one  time.  Latham  (1949)  gives 
an  extensive  account  of  the  species’  distribution  and  ecology  on  the 
island.  I  have  only  seen  C.  alpestris  twice  on  Long  Island.  The  first 
observation  was  in  a  pine  barren  area  south  of  Riverhead  and  was  repre¬ 
sented  by  a  tiny  fragment  of  a  thallus,  possibly  blown  there  from  a  larger 
colony  nearby  which  I  could  not  locate.  Latham  took  me  to  his  “Colony 
seven”  (Latham,  1949)  at  Napeague  Beach  which,  at  one  time,  was 
“in  excess  of  300  feet  in  diameter”  but  at  the  time  of  our  visit  consisted 
of  but  a  few  plants  scattered  among  low  shrubs,  bearberry  and  Cladonia 
submitis. 

Distribution  —  (Figure  20)  Arctic-boreal  element,  circumboreal 
(Ahti,  1961). 

Section  TENUES  Abb. 

129.  Cladonia  subtenuis  (Abb.)  Evans.  Trans.  Conn.  Acad.  Arts 
Sci.  35:  536.  1944.  Cladonia  tennis  *  Cl.  subtenuis  Abb.  Bull.  Soc.  Sci: 
Bretagne  16:  108.  1939. 

f.  subtenuis 

Material  seen  —  KINGS  COUNTY:  Forest  Park,  Hulst,  Novem¬ 
ber  31  (sic),  1890  (BKL  031993).  NASSAU  COUNTY:  Valley  Stream, 
Warner,  November  17,  1900  (BKL).  SUFFOLK  COUNTY:  95  speci¬ 
mens  collected  by  Imshaug  and/or  Brodo;  92  specimens  collected  by 
Latham  (Latham);  East  Point,  Taylor  32,  July  2  to  3,  1918  (BKL); 
Orient  Point,  Dillman,  1927  (NY);  Calverton,  Latham  7869,  Septem¬ 
ber  17,  1937  (NYS);  10  specimens  collected  by  R.  H.  Torrey  (NY); 
1.3  miles  W  of  Middle  Island,  S.  Smith  17715,  March  12,  1955  (NYS); 
Eastport  (vicinity),  S.  Smith  28512,  28511,  28510,  August  5,  1959 
(NYS). 

f.  cinerea  Ahti,  Ann.  Bot.  Soc.  [Zool.  Bot.  Fenn.]  ‘Vanamo’  32(1):  69. 
1961. 

Material  seen  —  SUFFOLK  COUNTY :  Promised  Land,  Latham 
27630,  June  2,  1951  (Latham)  (Holotype);  Peconic,  Latham  23445, 
April  11,  1945  (Latham). 

Ahti  (1961)  presents  a  full  discussion  of  f.  cinerea,  which  differs 
from  f.  subtenuis  only  in  lacking  usnic  acid. 


200 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


If  the  thalli  of  C.  subtemiis  are  fertile,  which  is  rare,  the  branches 
are  shorter,  stouter,  and  more  verrucose  than  sterile  specimens. 

Cladonia  subtemiis  and  C.  arbuscula  are  the  two  species  in  the  Long 
Island  Cladinae  most  difficult  to  separate.  They  have  the  same  chemical 
constituents  ( fumarprotocetraric  acid,  usnic  acid,  and  ursolic  acid)  and 
their  morphologies  overlap  to  a  large  degree. 

Ahti  (1961)  separates  the  two  largely  as  is  shown  in  the  following 
table: 

C.  subtemiis  C.  arbuscula 


1.  branching  mostly  dichotomous 

2.  axils  mostly  closed 

3.  slender  branches 

4.  main  branch  often  sub-  to 
indistinct 

5.  branchlets  mostly  erect 

6.  podetial  surface  smooth 

7.  pycnidial  jelly  red 


1 .  branching  mostly  tri-  and 
tetrachotomous 

2.  axils  mostly  open 

3.  heavy,  robust  branches 

4.  main  branch  robust  and  very 
distinct 

5.  branchlets  mostly  unilaterally 
falcate 

6.  podetial  surface  ±  warty 

7.  pycnidial  jelly  hyaline 


Upon  examining  specimens  determined  by  Ahti  as  C.  subtemiis  and 
C.  arbuscula,  and  after  personally  examining  scores  of  specimens  from 
Long  Island  and  nearby  Cape  Cod  and  southern  New  Jersey,  it  appears 
that  only  a  few  of  these  characters  approach  constancy. 

There  are  many  specimens  of  C.  subtemiis  which  are  quite  robust 
and  have  heavy  main  stems,  unlike  typical  subtemiis.  On  occasional  speci¬ 
mens,  axils  may  commonly  be  open  and  sometimes  may  even  show 
whorls  of  branchlets  around  the  gaping  hole,  although  this  latter  condition 
is  very  rare.  The  pycnidial  jelly  of  specimens  so  closely  approaching  C. 
arbuscula  should  be  examined.  The  jelly  will  be  reddish  or  red-brown 
in  subtemiis  and  colorless  in  arbuscula.  This  was  done  in  some  of  the 
questionable,  very  robust  specimens  of  subtemiis  from  Long  Island,  and 
helped  establish  the  range  of  variation  to  be  expected  in  this  very  vari¬ 
able  species.  Unfortunately,  pycnidial  jelly  can  only  be  examined  from 
relatively  fresh  specimens  (not  more  than  a  few  years  old). 

Cladonia  subtemiis  is  most  characteristic  of  partially  shaded  oak  or 
pine-oak  forests  but  can  also  be  found  in  open  sand  barrens  associated 
with  C.  submitis  and  C.  boryi. 

Distribution  —  (/.  subtemiis)  —  Eastern  United  States  (map:  Ahti, 
1961):  Temperate  e'ement.  East  Temperate  subelement;  British  Guiana 
(map:  Ahti,  1961). 

(/.  cinerea)  —  New  England  (Ahti,  1961). 

130.  Cladonia  terrae-novae  Ahti,  Ann.  Bot.  Soc.  ‘Vanamo’  30(4): 
11.  1959  (nomen  nudum);  Arch.  Soc.  ‘Vanamo’  14:  131.  1960. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk,  Latham  38016. 
August  19,  1963  (Latham). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


201 


The  rarity  of  this  species  on  Long  Island  is  difficult  to  understand 
in  view  of  its  abundance  on  Nantucket  Island  and  Cape  Cod  and  its 
presence  in  southern  New  Jersey.  In  all  but  a  few  cases  it  was  collected 
in  boggy  heath  or  on  bog  hummocks. 

The  presence  of  atranorin  along  with  usnic  acid  is  usually  sufficient 
to  separate  this  species  from  all  other  Cladinae  with  anisotomic  branching. 
Perlatolic  acid  is  also  present,  but  is  sometimes  difficult  to  demon¬ 
strate.  Although  the  species  shows  considerable  morphological  variability, 
especially  in  the  thickness  of  the  podetia  and  density  of  the  young 
branches,  the  rough  “tomentose”  appearance  of  the  surface  and  very 
discontinuous  algal  layer  is  striking  and  characteristic.  The  thallus  is 
always  distinctly  anisotomic  (having  one  or  more  distinct  main  stems) 
and  in  this  respect  differs  from  the  very  rare  usnic  acid  form  of  Cladonia 
evansii  which  has  a  similar  thallus  surface  and  also  contains  atranorin 
(p.  198). 

Cladonia  terrae-novae  differs  from  its  vicarious  European  parent 
species,  C.  impexa  Harm.,  principally  in  the  absence  of  atranorin  in  the 
latter,  although  Ahti  (1961)  asserts  that  there  are  good  morphological 
differences  between  the  two  species  as  well. 

An  interesting  phytogeographical  parallel  with  the  distribution  of 
C.  terrae-novae  involves  the  tiny  fern,  Schizaea  pusilla.  The  two  species 
have  their  southernmost  locality  in  the  very  same  bog  in  southern  New 
Jersey  and  are  found  growing  there  almost  side  by  side.  The  fern  has  not  yet 
been  found  on  Long  Island,  but,  like  the  lichen,  reoccurs  farther  north 
(especially  in  Nova  Scotia  and  Newfoundland)  in  great  abundance. 

Distribution  —  Along  the  Atlantic  coast  from  Newfoundland  to  New 
Jersey  (map:  Ahti,  1961):  Temperate  element,  Oceanic  subelement. 

Section  CLADINA 

131.  Cladonia  rangiferina  (L.)  G.  Web.  in  Wigg.  Prim.  FI.  Hol- 
saticae  90.  1780.  Lichen  rangiferinus  L.  Sp.  PI.  1153.  1753. 
subsp.  rangiferina  var.  rangiferina. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1018  (82),  1 447 
(83);  25  specimens  collected  by  Latham  (Latham). 

Cladonia  rangiferina ,  like  C.  alpestris,  is  a  rare  member  of  the  com¬ 
munity  on  open  sand-dunes  and  sand  plains  on  Long  Island,  but  is 
found  abundantly  in  the  Cape  Cod  region  in  the  same  community. 
Both  species  were  previously  more  common  on  the  island  than  they 
are  now  (p.  276). 

Distribution  —  Throughout  arctic,  boreal,  east  temperate,  and  west 
montane  North  America  (map:  Ahti,  1961):  Arctic-boreal  element; 
circumboreal. 

132.  Cladonia  arbuscula  (Wallr.)  Rabenh.  Deutschl.  Kryp.  FI.  2: 
110.  1845.  Patellaria  foliacea  var.  m.  Arbuscula  Wallr.  FI.  Crypt.  Germ. 
1:  425.  1831. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-284  (82),  3242 
(120);  Calverton,  Latham  7547,  October  10,  1933  (Latham);  Flanders, 


202  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Latham  24717  (Latham);  Calverton,  R.  H.  Torrey  (with  C.  subtenuis) , 
1936  (NY). 

The  nomenclatural  problems  of  this  species,  called  C.  sylvatica  (L.) 
HofFm.  by  most  authors  are  discussed  in  detail  by  Ahti  (1961). 

The  Long  Island  material  of  this  species  belongs  to  Ahti’s  subsp. 
arbuscula,  chemical  strain  I  (with  fumarprotocetraric  acid).  The  similari¬ 
ties  between  C.  arbuscula  and  C.  subtenuis  are  discussed  under  the  latter 
species  (p.  200). 

Cladonia  arbuscula  was  found  associated  with  C.  submitis  and 
Cetraria  islandica  on  sand  dunes. 

Distribution  —  (maps:  Ahti,  1961).  (sens,  lat.)  —  Arctic-boreal 
element;  circumboreal.  (subsp.  arbuscula,  chemical  strain  I)  —  Eastern 
boreal  and  temperate  North  America;  Temperate  element.  North  Tem¬ 
perate  subelement  (?),  but  clearly  boreal  to  arctic  in  Eurasia. 

133.  Cladonia  submitis  Evans,  Rhodora  45:  435.  1943. 

Material  seen — KINGS  COUNTY:  Forest  Park,  Hulst,  November  31 
(sic),  1890  (BKL  031993).  SUFFOLK  COUNTY:  66  specimens  collected 
by  Imshaug  and/or  Brodo;  36  specimens  collected  by  Latham  (Latham); 
Southampton,  Clute,  September  3-7,  1898  (NY);  East  Point,  Taylor  32, 
July  2-3,  1918  (BKL);  Farmingville,  Davis,  August  1916  (Staten  Island); 
Pike’s  Beach,  Westhampton,  R.  H.  Torrey ,  1936  (NY);  Holtsville,  R.  H. 
Torrey,  1936,  (NY);  Selden,  R.  H.  Torrey,  1936  (NY);  Route  112 
north  of  Coram,  R.  H.  Torrey,  1936  (NY);  Reeves  Bay  near  Flanders, 
R.  H.  Torrey,  1937  (NY);  Riverhead,  S.  Smith  28444,  28559,  28560, 
August  7,  1959  (NYS);  Selden,  S.  Cain  353,  June  30,  1936  (NY);  Noy- 
ack,  Latham  26423,  April  17,  1947  (FH). 

Ahti  (1961)  reported  that  an  isotype  specimen  of  C.  submitis  that 
he  examined  appeared  to  be  C.  mitis.  Upon  checking  the  holotype  speci¬ 
men  in  the  Evans  herbarium  I  discovered  that  it  was  already  annotated 
by  Ahti  (in  1961 )  and  by  Thomson  (in  1962).  Ahti  marked  it  as  pseudo- 
norangiformic  absent,  but  Thomson  noted  that  with  the  help  of  Kurokawa, 
he  found  pseudonorangiformic  acid  in  small  amounts  in  the  greater  por¬ 
tion  of  the  material.  I  attempted  to  recrystallize  the  crucial  chemical 
myself,  but  met  with  no  success.  In  this  connection,  it  should  be  men¬ 
tioned  that  the  holotype  material  is  not  a  typical  example  of  C.  submitis 
from  a  morphological  point  of  view.  Although  some  branchlets  show  the 
characteristic  prongs  and  robust  nature  of  the  species,  most  of  the 
material  is  rather  slender.  The  Sandstede  exsiccats  nos.  1564  and  1565, 
both  on  the  same  sheet  as  the  holotype  and  both  annotated  by  Evans  as 
being  C.  submitis  and  containing  “C”  (pseudonorangiformic  acid),  are 
much  more  representative  of  typical  C.  submitis. 

Cladonia  submitis  seems  to  have  two  basic  growth  forms  on  Long 
Island:  one  is  prostrate  and  sprawling  and  the  other  is  erect  and  often 
tufted.  The  former  is  characteristc  of  the  isolated  thalli  in  open  sand 
dune  areas  and  exposed  sand  barrens;  the  upright  form  is  usually  seen 
in  protected  situations,  between  clumps  of  grass,  in  extensive  colonies 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  203 

on  the  dunes,  and  in  partially  shaded  localities.  This  latter  form  often 
appears  very  much  like  C.  arbuscula,  which,  however,  is  PD  +  red  and 
lacks  pseudonorangiformic  acid.  The  prostrate  form  has  no  parallel  in 
the  Cladinae  and  is  easily  identified  in  the  field. 

It  is  interesting,  although  puzzling,  that  C .  submitis  is  very  abundant 
in  south  shore  dune  habitats,  but  is  entirely  absent  from  very  similar 
habitats  on  the  north  shore  (figure  75).  There  are  three  observable 
factors  correlating  with  its  occurrence  on  the  south  shore:  the  presence 
of  a  continuous  foredune  between  the  community  and  ocean,  the  pres¬ 
ence  of  Pinus  rigida  in  the  immediate  areas,  and  the  high  acidity  of  the 
sand.  These  three  factors  are  probably  somewhat  interrelated  and  may 
affect  the  Cladonia  distribution  directly  or  indirectly  and  act  either  inde¬ 
pendently  or  together. 

It  is  known  that  salt  mist  and  salt  spray  causes  maritime  substrates 
to  become  more  alkaline  than  normal  ( Barkman,  1958).  It  is  also 
known  that  Pinus  rigida  is  intolerant  of  large  quantities  of  salt  spray 
(Boyce,  1954).  Ahti  (1961)  stated  that  Cladonia  submitis  is  intolerant  to 
salt  spray  and  is  never  found  near  the  ocean. 

All  these  facts  seem  to  suggest  strongly  that  the  salt  spray  on  the 
south  shore,  blocked  to  a  large  extent  by  the  foredune,  is  not  nearly  as 
abundant  as  it  is  on  the  north  shore  where  the  only  protection  comes 
from  low  dunes  and  hollows  (see  Oostings  and  Billings,  1942).  It  is, 
therefore,  the  salt  spray,  rather  than  any  directly  observed  factor  such 
as  sand  pH  or  the  accumulation  of  pine  detritus,  which  very  likely  limits 
the  distribution  of  C.  submitis. 

Distribution  —  Atlantic  coastal  plain  (map:  Ahti,  1961 ) :  Temperate 
element,  Coastal  Plain  subelement;  Japan  (Ahti,  1961). 

134.  Cladonia  mitis  Sandst.  Clad.  exs.  no.  55.  1918. 

Material  seen  —  SUFFOLK  COUNTY :  Orient,  West  Long  Beach, 
Latham  23437,  April  7,  1945  (Latham);  Promised  Land,  Latham  25473, 
April  1,  1946  (Latham). 

This  species  is  extremely  rare  on  Long  Island.  It  is  apparently  a 
member  of  the  community  on  exposed  sand  with  other  Cladinae. 

Distribution  —  Throughout  the  arctic  and  boreal  northern  hemi¬ 
sphere  (map:  Ahti,  1961):  Arctic-boreal  element;  circumboreal. 

UMBIUCARIACEAE 

36.  UMBILICARIA  Hoffm. 

135.  Umbilicaria  mammulata  (Ach.)  Tuck.  Proc.  Amer.  Acad.  Arts 
Sci.  1:  261.  1847.  Gyrophora  mammulata  Ach.  Syn.  Lich.  67.  1814. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3843  (76);  Mon- 
tauk,  Latham,  May  1920  (Latham);  Plum  Island,  Latham,  July  1931 
( Latham ) . 

Llano  (1950)  considered  Tuckerman’s  transfer  of  Gyrophora  mam¬ 
mulata  Ach.  to  Umbilicaria  invalid,  since  Tuckerman,  not  knowing  the 
true  identity  of  Acharius’  species,  was  actually  working  with  what  is  now 


204  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

known  as  U.  caroliniana  Tuck.  Llano,  therefore,  proposed  the  new  com¬ 
bination  U.  mammulata  (Ach.)  Llano  and  considered  U.  mammulata 
(Ach.)  Tuck,  non  Llano  as  a  synonym  of  U.  caroliniana.  Llano’s 
transfer  is  not  necessary,  since,  although  Tuckerman  was  mistaken  about 
the  identity  of  his  new  combination,  it  was  still  validly  published. 

The  species  has  often  been  considered  under  the  names  Umbilicaria 
dillenii  Tuck,  or  Gyrophora  dillenii  (Tuck.)  MiilL  Arg. 

Of  the  three  Umbilicariae  on  Long  Island,  this  species  is  the  only 
one  I  saw  growing  in  the  field.  It  was  found  on  exposed  granitic  boulders 
at  the  summit  of  a  morainal  hill  south  of  Riverhead.  Torrey  (1933)  also 
reported  it  from  the  Wading  River  region. 

Distribution  —  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit;  endemic  (map:  Llano,  1950). 

136.  Umbilicaria  muhlenbergii  (Ach.)  Tuck.  Enum.  N.  Amer.  Lich. 
55.  1845.  Gyrophora  muhlenbergii  Ach.  Lich.  Univ.  227.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Gardiner’s  Island,  Latham, 
June  28,  1927  (Latham);  Bald  Hill,  3  miles  S.  of  Calverton,  Latham , 
July  1,  1937  (Latham);  Yaphank,  Wm.  Davis,  January  3,  1929  (Staten 
Island). 

This  species  is  treated  in  the  genus  Actinogyra  by  Llano  (1950).  It 
is  found  on  boulders. 

Distribution  —  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit;  Europe  (Poelt,  1963);  north  temperate  regions 
of  Asia  (map:  Llano,  1950). 

137.  Umbilicaria  papulosa  (Ach.)  Nyl.  Mem.  Soc.  Sci.  Nat.  Cherb. 
5:  107.  1857.  Gyrophora  papulosa  Ach.  Lich.  Univ.  226.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Wading  River,  Latham 
2643.  July  20,  1926  (Latham). 

Llano  (1950)  considered  this  species  in  the  genus  Lasallia. 

The  species  apparently  is  unknown  on  Long  Island  outside  the 
Wading  River  region.  I  have  searched  the  area  for  Umbilicariae  without 
success,  but  Latham  (see  above)  and  Raymond  Torrey  (1933)  collected 
U.  papulosa  there. 

On  Cape  Cod  ( Barnstable  County,  N  of  Woods  Hole,  Brodo  3927, 
3956),  I  collected  several  specimens  of  this  species.  It  was  growing 
abundantly  over  almost  all  exposed  and  partially  shaded  boulders  in 
the  area,  but  was  found  nowhere  else  on  the  Cape. 

Distribution  —  Temperate  e'ement,  Appalachian  subelement.  Appa¬ 
lachian-Great  Lakes-Rocky  Mountain  unit,  with  several  west  coast  locali¬ 
ties;  Africa  (map:  Llano,  1950). 

ACAROSPORACEAE 

37.  SARCOGYNE  Flot. 

138.  Sarcogyne  clavus  (Ram.  in  Lam.  &  DC.)  Kremp.  Denkschr. 
Kgl.  Bayer.  Bot.  Ges.  4:  212.  1861.  Lichen  clavus  Ram.  in  Lam.  &  DC. 
FI.  Franc,  ed.  3,  2:  348.  1805. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  205 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  779  (90A),  786 
(90A),  1715  (133),  1803  (127),  2383  (123),  2705  (111),  2810  (106), 
3354  (62),  3377  (94),  3306  (134),  3432  (134),  3848  (76);  Orient, 
Latham  22246,  May  3,  1914  (Latham);  probably  Montauk  Point,  von 
Scheur,  July  22,  1895  (MO). 

Fink  (  1935)  treats  this  species  in  the  genus  Biatorella. 

Sarcogyne  davits  is  found  on  exposed  granitic  boulders. 

Distribution  —  Connecticut,  New  York,  Alabama,  and  California 
(Magnusson,  1935);  Minnesota,  Black  Hills:  Temperate  element,  North 
Temperate  subelement  (?);  Europe. 

139.  Sarcogyne  privigna  (Ach.)  Mass.  Geneac.  Lich.  10.  1854. 
Lecidea  privigna  Ach.  Meth.  Lich.  49.  1803. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  961  (S  of  50). 

The  similarity  of  this  species  with  S.  clavus  and  S.  simplex  (as  well 
as  S.  pruinosa )  is  discussed  in  some  detail  by  Magnusson  (1935). 
Sarcogyne  privigna  is  similar  to  S.  clavus  in  having  a  smooth,  red-black 
apothecial  disk,  but  differs  from  the  latter  in  having  small  (less  than 
1  mm  broad),  concave,  irregular  apothecia  with  prominent  margins. 

Distribution  —  New  Hampshire,  Connecticut,  and  New  Mexico 
(Magnusson,  1935);  Black  Hills;  Europe. 

140.  Sarcogyne  simplex  (Dav.)  Nyl.  Mem.  Soc.  Sci.  Nat.  2:  337. 
1854.  Lichen  simplex  Dav.  Trans.  Linn.  Soc.  Lond.  2:  283.  1793. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1907  (114),  3089 
(128),  3255  (119);  Orient,  Latham,  April  25,  1921  (Latham). 

This  species  is  the  only  Sarcogyne  with  small  apothecia  having  rough 
disks.  It  was  collected  on  exposed  or  partially  shaded  granitic  boulders. 

Distribution  —  Maine,  Connecticut,  Tennessee,  Minnesota,  Black 
Hills,  Washington,  Manitoba;  Canadian  archipelago  (Thomson,  1960): 
Arctic-boreal  element  (?);  circumboreal. 

38.  ACAROSPORA  Mass. 

141.  Acarospora  fuscata  (Schrad.)  Arn.  Verhandl.  Zool.  -Bot.  Ges. 
Wien  20:  528.  1870.  Lichen  fuscatus  Schrad.  Spicil.  FI.  Germ.  83.  1794. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3513  (10).  SUF¬ 
FOLK  COUNTY:  Imshaug  25561  (52);  Brodo  1556  (103),  1739 
(126),  2373  (123),  2660  (108),  2736  (111),  3386  (94),  3449  (134), 
3849  (76),  3883  (62);  16  specimens  collected  by  Latham  (Latham); 
Montauk,  Hither  Beach,  Latham  27289,  October  28,  1947  (MO). 

Both  Magnusson  (1929)  and  Weber  (1962)  have  commented  on 
the  extreme  morphological  variability  of  this  species.  The  C  +  red  re¬ 
action  of  the  cortex  is  also  somewhat  variable,  being  strongly  positive  in 
some  cases  and  almost  negative  in  others. 

Acarospora  fuscata  is  found  on  granite  boulders  and  pebbles  in 
exposed  or  partially  shaded  localities  (figure  57).  In  addition,  one 
questionable  specimen  was  collected  on  calcareous  rock  ( Latham 
22332,  with  Lecanora  dispersa),  and  one  was  found  growing  on  a 
storm-tide-washed  boulder  (hygrohaline  zone)  ( Brodo  2736). 


206 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Distribution  — -  Northern  and  middle  states  (Fink,  1935):  Tem¬ 
perate  element,  North  Temperate  subelement;  Europe;  Asia  ( Mag- 
nusson,  1929). 


PERTUSARIACEAE 

39.  PERTUSAR1A  DC. 

142.  Pertusaria  alpina  Hepp  in  Ahles,  Pertus.  et  Conotr.  12.  1860. 

Material  seen  —  SUFFOLK  COUNTY:  Orient,  Latham,  May  23, 

1914  (Latham). 

The  Long  Island  specimen  was  compared  with  Cummings’  exsic- 
cats  (Decades  of  North  American  Lichens  no.  281  and  Lichens  Boreali- 
Americani  no.  211),  the  former  of  which  was  cited  by  Erichsen  (1941) 
as  P.  alpina.  (These  exsiccats  are  both  mixtures  of  P.  pustulata,  which 
has  2  spores  per  ascus,  and  P.  alpina  which  has  4  to  8  spores.)  The 
Long  Island  specimens  were  morphologically  and  anatomically  identical 
with  these  exsiccats,  but  the  Cummings  specimens  contained  stictic  acid 
(by  chromatography)  and  were  K  +  yellow  and  PD  +  orange,  whereas 
the  Long  Island  specimen  was  K  —  or  K  +  yellowish  and  PD  — .  (The 
specimen  was  too  scanty  to  extract  for  chromatography.)  The  ultra¬ 
violet  fluorescence  of  the  thallus  (orange-pink)  was  the  same  in  all  the 
material,  however. 

The  Latham  specimen  is  on  cedar  lignum  and  not  on  bark,  as  is  the 
case  with  the  Cummings  material. 

Distribution  —  Nova  Scotia,  District  of  Columbia  (Cummings’  Dec¬ 
ades  no.  281),  Michigan;  Europe. 

143.  Pertusaria  amara  (Ach.)  Nyl.  Bull.  Soc.  Linn.  Norm.  II. 
6:  288.  1872.  Variolaria  amara  Ach.  Kgl.  Vet.  -Akad.  Nya  Hand!.  163. 
1809. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  890  (56),  902  (56), 
1411  (83),  1417a  (83),  2806  (102),  3898  (112);  Orient,  Long  Beach, 
Latham  22338,  December  7,  1944  (Latham);  Orient,  Latham  61,  May 
10,  1914  (Latham). 

This  species  is  the  only  KC  +  violet  Pertusaria  on  Long  Island. 
It  was  always  found  sterile.  Pertusaria  amara  grows  on  the  bark  of  vari¬ 
ous  trees,  usually  in  or  near  bogs  (figure  38). 

Distribution  —  Nova  Scotia,  Quebec,  Maine,  Connecticut,  the  Smoky 
Mountains  of  Tennessee,  North  Carolina,  Michigan,  Wisconsin,  Black 
Hills;  Washington  (Fink,  1935):  Temperate  element.  North  Temperate 
subelement;  Europe. 

144.  Pertusaria  multipuncta  (Turn.)  Nyl.  Lich.  Scand.  179.  1861. 
Variolaria  multipuncta  Turn.  Trans.  Linn.  Soc.  Lond.  9:  137.  1808. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  857  (47);  Greenport, 
Latham  1983,  February  27,  1923  (Latham);  Montauk,  Latham  3944 
(sterile),  April  7,  1927  (Latham):  Greenport,  Latham  27287,  April  16, 
1945  (Latham);  Greenport,  Latham,  February  27,  1923  (Latham). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


207 


The  Pertusaria  multipuncta  group  seems  to  be  a  rather  heterogeneous 
complex  of  KOH  — ,  PD  —  taxa  having  one  spore  per  ascus,  and  in¬ 
cludes  at  least  three  populations  having  spore  size  ranges  which  do  not 
overlap.  Representatives  of  two  of  these  populations  are  on  Long  Island 
and  seem  to  be  morphologically  distinct  as  well  (see  key).  This  group 
certainly  needs  further  study. 

These  specimens  were  found  on  the  bark  of  various  deciduous 
trees. 

Distribution  —  No  comment  seems  warranted  until  the  taxonomy 
of  the  group  is  clarified. 

145.  Pertusaria  propinqua  Mull.  Arg.  Flora  67:  273.  1884. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3276  (119). 

This  species,  though  represented  by  only  a  single  collection  from 

Long  Island,  was  found  abundantly  in  the  locality  where  it  occurred. 

The  description  of  P.  rubescens  Erichs.  (Erichsen,  1941)  agrees  very 
well  with  the  Long  Island  material  except  for  the  lack  of  zoned  spore 
walls  in  the  former.  The  type  specimen  of  P.  rubescens  is  from  a  hickory 
in  the  New  Jersey  coastal  town  of  “Sea  Girton”  (Sea  Girt?).  This 
exactly  parallels  the  Long  Island  collection  in  a  coastal  oak-hickory  woods 
on  Carya  cfr.  tomentosa.  Without  having  seen  authentic  material  of 
P.  rubescens,  I  am  not  listing  it  as  a  synonym. 

Although  P.  propinqua  was  described  from  a  specimen  on  granite,  it 
appears  to  be  identical  with  the  original  collection  of  P.  torquata  Miill. 
Arg.,  which  was  on  bark.  I  saw  the  original  material  of  both  taxa  (in 
herb.  MICH).  Since  the  species  is  poorly  known,  a  short  description 
of  the  Long  Island  material  follows: 

Thallus  dark  ashy  grey,  smooth  to  rugose,  becoming  thick  and 
cracked;  fruit  warts  smooth  or  rough,  becoming  distinctly  constricted 
at  the  base  in  maturity,  lighter  in  color  than  the  thallus  (appearing  as  if 
their  top  surfaces  were  rubbed),  1-2  mm  in  diameter;  ostioles  single  to 
many,  usually  large,  ashy  to  black,  usually  somewhat  depressed;  epi- 
thecium  cinereous,  turning  violet  in  KOH;  spores  8,  irregularly  arranged 
in  the  ascus,  89-96  x  40-41  (j.;  spore  walls  zoned.  Medulla  of  fruit  warts 
and  thallus  PD  +  yellow,  KOH  +  deep  blood  red.  Norstictic  acid  dem¬ 
onstrated  in  KOH. 

Distribution  —  Temperate  element.  Coastal  Plain  subelement  (Fink, 
1935  sub  P.  marginata  Nyl.);  endemic. 

146.  Pertusaria  subpertusa  sp.  nov. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1 035  (112),  1436 
(83),  1674  (88),  2163  (102),  2289  (87). 

Thallus  virido-cinerous,  continuus,  rimae  acquirendus,  rugosus, 
ultimus  minutissime  verrucosus;  verrucae  fructae  plerumque  dis¬ 
perses,  (0.5-)0.65-l .10  (-1 .30)  mm  diain.,  leves,  hemisphericales  ad 
subplaniferes;  colorae  thallis,  basibus  perspicues  constrictis;  ostioles 
3-7  per  verruca  fructus,  obscures  vel  pallides,  tantum  depressiuscules, 
0.05  ad  0.15  nun  diam.  Paraphyses  tenuisimae  (±  1  ij.)  ramiosissimae. 


208 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Asci  193-287  x  35-42  ij„  parietis  crcissa.  Sporae  hyalinae,  non 
septatae,  101-144(-161 )  x  35-45  p,,  parietes  4-10  p  crassae,  zonates  et 
canaliculates,  2  vel  rarissime,  l  per  ascus.  Epithecium  obscurum, 
KOH  +  violaceum.  Medulla  verrucarum  fructus  et  thalli  PD  + 
rubro-aurantiaca,  KOH  +  luteus  transeuns  rubidus,  C  — ,  KC  — . 
Materiae  chemicae:  acidmn  fiunarprotocetraricum  et  acidum  sala- 
cinicum.  Corticola. 

Holotype:  SUFFOLK  COUNTY:  Three  Mile  Harbor,  on  Old  North¬ 
west  Road  0.7  miles  from  junction  with  Alewife  Brook  Road,  Brodo 
1035,  July  12,  1960,  on  bark  of  Acer  rubrum  in  bog  (MSC)  (figures  84, 
85). 


Figure  84.  Pertusaria  subpertusa  (holotype).  Scale  equals  1  mm.  Draw¬ 
ing  by  Brenda  Carter  Haas. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


209 


Figure  85.  Pertusaria  subpertusa  (holotype).  Spores  and  ascus  mounted 
in  water.  Section  of  fruiting  wart  shows  two  apothecia,  one  in  median 
view.  Fine  stippled  shading  indicates  algal  layer;  coarse  stippling 
indicates  dark  to  opaque  areas  in  the  fungal  tissue.  Drawings  made 
with  the  aid  of  a  camera  lucida  apparatus. 


210  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Thallus  grey  to  greenish  grey,  continuous,  becoming  cracked,  rugose, 
and  finally  minutely  verrucose;  fruit  warts  mostly  scattered,  (0.5-)0.65 
-1.10  (-1.30)  mm  in  diameter,  smooth,  hemispherical  to  flattened,  the 
same  color  as  the  thallus,  distinctly  constricted  at  the  base;  ostioles  3  to 
7  per  fruit  wart,  dark  or  pale,  only  slightly  depressed,  0.05-0.15  mm  in 
diameter.  Paraphyses  very  slender  (ca.  Ijj.),  much  branched;  asci  thick- 
walled,  193-287  x  36-45  [jl;  spores  hyaline,  nonseptate,  101-1 44 (-161 )  x 
35-45  [x,  walls  4-10  p.  thick,  zoned  and  channelled,  2  or,  very  rarely,  1 
spore  per  ascus;  epithecium  dark,  KOH  +  violet.  Medulla  of  fruit  warts 
and  thallus  PD  +  red-orange,  KOH  +  yellow,  becoming  dark  red,  C  — , 
KC  — ;  fumarprotocetraric  acid  present.  Corticolous. 

The  Long  Island  material  is  rather  uniform  in  morphology,  but  does 
show  some  variation  in  the  color  of  the  ostioles  (becoming  pale  in  some 
specimens)  and  in  the  depth  and  breadth  of  the  ostiole  depressions  (often 
becoming  very  deep  and  up  to  0.20mm  broad  in  maturity). 

The  epithet  “subpertusa”  is  used  for  this  new  species  to  emphasize 
its  similarity  in  general  appearance  and  spore  type  to  P.  pertusa  (L. ) 
Tuck.  Pertusaria  pertusa  has  larger  spores  (145-229  x  40-82  jj.)  and  con¬ 
tains  stictic  acid.  All  other  North  American  KOH  +  red  Pertusariae 
have  norstictic  acid  rather  than  salacinic  acid  and  none,  to  my  knowledge, 
contains  fumarprotocetraric  acid  as  well. 

Of  the  four  Long  Island  specimens,  three  were  found  growing  on  the 
bark  of  Acer  rubrum  in  bogs  or  swamps  and  one  was  on  Quercus  velutina 
bark.  I  also  collected  a  specimen  in  southern  New  Jersey  (Atsion,  Brodo 
3587)  on  a  black  oak  just  outside  a  bog. 

Distribution  —  New  Jersey;  endemic. 

147.  Pertusaria  trachythallina  Erichs,  in  Degel.  Ark.  Bot.  30A(  1 ) : 
36.  1940. 

Material  seen  —  SUFFOLK  COUNTY:  31  specimens  collected  by 
Imshaug  and/or  Brodo. 

This  species  is  discussed  at  length  by  Erichsen  in  his  original  descrip¬ 
tion. 

The  species  is  found  on  the  bark  of  various  deciduous  trees,  usually 
at  breast  height.  It  can  be  considered  a  member  of  the  black  oak,  breast 
height  community,  although  it  has  also  been  found  on  Quercus  alba  and 
Fag  us  grandifolia. 

Distribution  —  Maine;  endemic. 

148.  Pertusaria  tuberculifera  Nyl.  Act.  Soc.  Scien.  Fenn.  7:  448. 
1863. 

Material  seen  —  QUEENS  COUNTY :  Ridgewood,  G .  B.  Brainerd, 
(1866?)  (BKL  031906);  Ridgewood,  G.  B.  Brainerd,  (1866?)  (BKL). 
SUFFOLK  COUNTY:  24  specimens  collected  by  Imshaug  and/or  Brodo; 
Montauk,  Latham  27286,  April  17,  1946  (Latham);  Springs,  East  Hamp¬ 
ton,  Latham  28321,  February  9,  1949  (Latham). 

Pertusaria  tuberculifera  belongs  to  Erichsen’s  subgenus  Eupertusaria 
section  Insensibiles.  The  material  treated  here  was  probably  considered 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  211 

under  the  name  P.  leioplaca  (Ach.)  Lam.  &  DC.  in  Fink  (1935)  where 
P.  leioplaca  is  described  as  having  4  to  8  spores.  Erichsen  (  1935)  regards 
P.  leioplaca  to  be  4-spored  alone,  or  rarely  2,  3,  or  5-spored.  The  Long 
Island  specimens  are  all  dominantly  8-spored,  with  the  4  spore  condition 
occurring  frequently  in  the  same  apothecia.  Pertusaria  tuberculifera  and 
P.  tetrathalamia  (Fee)  Nyl.  are  often  considered  to  be  conspecific,  hut 
Erichsen  ( 1936)  pointed  out  that  the  latter  has  only  four  spores  per  ascus. 

Since  the  species  apparently  is  fairly  common  and  yet  poorly  known, 
a  more  detailed  description  of  the  Long  Island  material  may  have 
some  value: 

Thallus  dark  ashy  grey,  continuous,  smooth,  becoming  rugose  and 
verrucose;  fruit  warts  large,  1-3  mm  in  diameter,  irregular,  crowded, 
distinctly  constricted  at  the  base  in  maturity;  spores  4  to  8,  hyaline, 
(30-)  34-40  x  ( 55-)  62-80 ( -97)  p,  walls  6  p  thick,  zoned,  smooth.  Medulla 
of  fruit  warts  KOH  — ,  PD  — ,  KC  — ,  C  — .  Thallus  UV  +  orange 
fluorescence. 

Distribution  —  South  America  (type  locality),  West  Indies  (Im- 
shaug,  1957b),  New  Jersey  (see  above):  Tropical  element,  Coastal 
Plain  subelement  (?). 

149.  Pertusaria  velata  (Turn.)  Nyl.  Lich.  Scand.  179.  1861.  Par- 
melia  velata  Turn.  Trans.  Linn.  Soc.  Lond.  9:  143.  1808. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1417B  (83),  2158 
(102),  2138  (102),  3101  (122),  3278  (119);  Riverhead,  Latham, 
May  16,  1960  (Latham);  Three  Mile  Harbor,  East  Hampton  Twp., 
Ogden  5405,  May  11,  1954  (NYS). 

The  C  +  red  disks  and  thallus  and  the  lecanorine  apothecia  of  this 
species  give  it  a  superficial  similarity  with  a  species  of  Ochrolechia.  How¬ 
ever,  the  very  large  spores,  one  per  ascus,  easily  refer  it  to  Pertusaria. 

Pertusaria  velata  is  usually  found  in  humid  forests  or  bogs  on  the 
bark  of  deciduous  trees.  Almborn  (1948)  stated  that  the  species  is 
typical  of  the  Pyrenula  nitida  society  on  Fagus  and  has  an  oceanic  affin¬ 
ity.  This  would  be  borne  out  to  some  extent  by  its  “oceanic”  distribution 
on  Long  Island. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Tennessee,  North 
Carolina,  Oklahoma,  Michigan,  Indiana,  Iowa,  Wisconsin,  Minnesota, 
Black  Hills,  Washington,  coastal  Alaska,  British  Columbia:  Temperate 
element,  Oceanic  subelement  (?);  Europe  (oceanic),  Asia,  Africa,  South 
America  (Almborn,  1948). 

150.  Pertusaria  xanthodes  Mull.  Arg.  Flora  67:  286.  1884. 

Material  seen  —  SUFFOLK  COUNTY :  80  specimens  collected  by 

Imshaug  and/or  Brodo;  10  specimens  collected  by  Latham  (Latham). 

The  Long  Island  material  agrees  well  with  Muller’s  type  specimen 
from  Texas,  which  I  saw  in  Geneva.  The  thallus  of  the  type  was  yellow, 
having  ampliariate  fruit  warts,  each  with  one  pale,  more  or  less  de¬ 
pressed  ostiole  and  containing  one  or  two  apothecia.  There  were  2  spores 


212 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


per  ascus  and  walls  were  clearly  zoned,  often  with  one  of  the  walls 
roughened  on  the  inner  surface. 

This  species  is  easily  confused  with  certain  forms  of  Pertusaria 
pustulcita  (Ach.)  Duby,  and  was  almost  surely  considered  under  this 
name  in  Fink  (1935).  Pertusaria  pustalata  is  characterized  by  spores 
with  thin,  smooth,  unzoned  walls,  and  by  dark  ostioles.  In  addition,  the 
epithecium  of  P.  pustalata  generally  turns  KOH  +  violet. 

Since  my  material  is  rather  variable,  I  will  break  down  its  descrip¬ 
tion  as  follows: 


Constant  characters: 

1 .  on  bark 

2.  spores  2  per  ascus 

3.  spores  30-45  x  70-120  w 

4.  spore  walls  zoned, 
thick,  rough 

5.  stictic  acid  present 

6.  UV  fluorescence  pink-orange 

Almost  constant  characters: 

1.  yellow  color  of  thallus 

2.  ampliariate  fruit  warts 

3.  hypophloedal  thallus 


Variable  characters: 

1.  ostiole  color  (pale  or  dark) 

2.  concentration  of  stictic  acid 

3.  thickness  of  thallus 

4.  degree  of  density  of 
fruit  warts 


Never  seen: 

1.  spore  walls  thin,  smooth 

2.  epithecium  KOH  +  violet 


Pertusaria  xanthodes  is  found  on  the  bark  of  various  species  of 
deciduous  trees,  usually  in  well-lighted  situations. 

Distribution  —  Cape  Cod  (Massachusetts),  New  Jersey,  Texas 
(type  locality),  West  Indies  (Imshaug,  1957b):  Temperate  element!?), 
Coastal  Plain  subelement;  endemic. 


40.  MELANARIA  Erichs. 

151.  Melanaria  macounii  Lamb,  Ann.  Rep.  Nat.  Mus.  Can.  132: 
286.  1954. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  903  (56),  1858 
(117),  2162  (102),  2804  (102),  3281  (119);  Napeague,  Latham 
2848 ,  March  1,  1927  (Latham). 

Melanaria  macounii  resembles  Pertusaria  pertusa  in  many  respects. 
Both  have  polycarpous,  smooth,  fruit  warts  of  the  same  color  as  the 
thallus,  both  have  two  spores  per  ascus,  and  of  approximately  the  same 
size  range,  and  both  contain  stictic  acid.  In  M.  macounii,  however,  the 
spores  are  distinctly  radiately  channeled  and  often  are  brownish,  turning 
sordid  violet  in  KOH.  (The  hyaline  spores,  which  predominate  in  the 
Long  Island  material,  do  not  give  this  KOH  reaction.) 

The  species  is  found  on  the  bark  of  various  deciduous  trees  in 
humid  forests  or  bogs. 

Distribution  —  Nova  Scotia  (type  locality);  Great  Lakes  region 
(seen  in  herb.  MSC);  endemic. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  213 

LECANORACEAE 

41.  IONASPIS  Th.  Fr. 

152.  lonaspis  odora  (Ach.  in  Schaer.)  Th.  Fr.  Lich.  Scand.  1:  273. 
1871.  Gyalecta  odora  Ach.  in  Schaer.  Lich.  Helv.  Spic.  2:  80.  1826. 

Material  seen  - —  SUFFOLK  COUNTY:  Shelter  Island,  Latham 
22881,  October  26,  1944  (Latham). 

Latham’s  specimen  agrees  perfectly  with  the  description  of  the 
species  given  by  Magnusson  (1933)  in  his  monograph  of  the  genus. 
Although  Tuckerman  (1882)  cited  a  specimen  of  I.  odora  from  New 
Hampshire,  Magnusson  (1933)  stated  that  the  species  is  “most  likely 
not  in  North  America.”  lonaspis  lavata  Magn.  was  described  from  a 
Merrill  collection  from  Mount  Rainier,  Washington.  This  species,  how¬ 
ever,  differs  from  /.  odora  in  having  confluent,  brownish  apothecia,  rather 
than  scattered  pale  pink  or  yellow-brown  apothecia.  For  some  unknown 
reason,  the  type  specimen  of  /.  lavata  is  absent  from  its  packet  (in  herb. 
FH)  and  I  therefore  have  not  seen  it. 

The  species  is  listed  under  Lecanora  by  Fink  (1935). 

In  addition  to  the  reports  of  this  species  from  the  White  Mountains 
of  New  Hampshire  (Tuckerman,  1882),  1  collected  a  specimen  from 
Cape  Cod,  Massachusetts  ( Brodo  4399B ),  on  a  granite  boulder  in  parti¬ 
ally  shaded  oak  woods,  a  similar  habitat  to  that  of  the  Long  Island 
collection.  Magnusson  (1933,  p.  20),  however,  states  its  ecology  as  “on 
granitic  stone  on  the  banks  in  brooks  and  lakes  at  least  part  of  the 
year  wetted  by  water.” 

Distribution  —  New  Hampshire  (Tuckerman,  1882),  Massachusetts 
(see  above);  Europe  (“boreal-alpine  species”)  (Magnusson,  1933). 

42.  LECANORA  Ach. 

153.  Lecanora  atra  (Huds.)  Ach.  Lich.  Univ.  344.  1810.  Lichen  ater 
Huds.  FI.  Angl.  445.  1762. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk,  Latham  24167, 
May  4,  1926  (Latham). 

The  species  is  usually  found  on  stone  or  tree  bark,  but  is  also  known 
to  occur  on  lignum  on  occasion  (Hillman  and  Grummann.  1957;  Erich- 
sen,  1957). 

Distribution  —  Alaska,  Washington,  Idaho,  Quebec,  Michigan,  Min¬ 
nesota,  Black  Hills,  Arizona:  Temperate  element  (?),  North  Temperate 
subelement,  reported  from  European  and  Asian  arctic  by  Lynge  (1938, 
1928). 

154.  Lecanora  caesiocinerea  Nyl.  in  Malbr.  Bull.  Soc.  Amis  Sci.  Nat. 
Rouen  5:  320.  1869. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3505 A  (10).  SUF¬ 
FOLK  COUNTY:  Brodo  3871  (62). 


214  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  Long  Island  specimens  of  L.  caesiocinerea  agree  well  with  Mag- 
nusson’s  description  of  this  species  (Magnusson,  1939)  except  that  they 
have  moniliform  rather  than  submoniliform  paraphyses  and  spores  which 
are  slightly  smaller  ( 1 5-20  [j,  long  rather  than  over  20  p.  long). 

This  species  is  very  similar  to  L.  cinerea  and  perhaps  should  be 
included  there.  It  dilfers  in  the  slightly  shorter  pycnoconidia,  the  KOH  — 
thalline  reaction,  and  the  fewer  apothecia.  (See  additional  notes  under 
L.  cinerea.) 

Lecanora  caesiocinerea  grows  on  exposed  or  partially  shaded  granitic 
boulders. 

Distribution  — -  Nantucket  Island  (Massachusetts)  (Brodo  4004), 
Black  Hills,  Arizona;  Europe. 

155.  Lecanora  caesiorubella  Ach.  Lich.  Univ.  366.  1810. 
subsp.  lathamii  Imsh.  &  Brodo,  Nova  Hedw.  12:24.  1966. 

Material  seen  —  NASSAU  COUNTY:  Brodo  559  (13).  SUFFOLK 
COUNTY:  103  specimens  collected  by  Imshaug  and/or  Brodo;  19  speci¬ 
mens  collected  by  Latham  (Latham);  Eastport,  Schrenk,  June  26,  1894 
(MO);  Greenport,  Latham  18.  August  2,  1914  (FH);  Greenport,  Latham 
195 ,  March  20,  1914  (FH). 

All  the  Long  Island  material  of  this  species  belongs  to  subspecies 
lathamii.  Since  a  complete  discussion  of  L.  caesiorubella  and  other  mem¬ 
bers  of  the  L.  pallida  group  has  recently  been  published  (Imshaug  and 
Brodo,  1966),  it  suffices  to  say  that  this  subspecies  of  L.  caesiorubella  is 
characterized  by  a  C  +  orange-yellow  disk  and  by  the  presence  of  pro- 
tocetraric  acid  and  norstictic  acid  in  apothecial  sections,  the  latter  being 
more  or  less  confined  to  the  stipe. 

This  subspecies  of  L.  caesiorubella  is  found  on  the  bark  of  decidu¬ 
ous  trees,  usually  in  exposed  areas. 

Distribution  (subspecies  lathamii)  —  Nova  Scotia  to  Texas:  Tem¬ 
perate  element.  Coastal  Plain  subelement;  endemic.  The  species  as  a 
whole  has  a  tropical-temperate  distribution  (including  South  America, 
Africa)  (Imshaug  and  Brodo,  1966). 

156.  Lecanora  chlarotera  Nyl.  Bull.  Soc.  Linn.  Norm.  II.  6:  274. 
1872. 

Material  seen  —  NASSAU  COUNTY:  Cold  Spring,  Grout.  April  1, 
1900  (BKL).  SUFFOLK  COUNTY:  83  specimens  collected  by  Imshaug 
and/or  Brodo;  14  specimens  collected  by  Latham  (Latham);  Montauk 
Point.  Easthampton  Twp.,  northeast  of  Prospect  Hill  between  Great  Pond 
and  Oyster  Pond,  Ogden  5411,  May  12,  1954  (MSC). 

This  species  is  very  common  on  Long  Island,  and  it  is  extremely 
variable.  The  color  of  the  disks,  for  examp'e,  varies  from  a  pale  yellow- 
brown  to  a  dark  chocolate  brown;  the  margins  are  usually  somewhat 
crenate,  but  sometimes  are  quite  smooth  and  even;  the  epithecium  is 
usually  conspicuously  granular  but  sometimes  is  almost  without  granules. 
Large,  irregular,  colorless  crystals,  however,  always  can  be  found  in  the 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  2  1  5 

amphithecium,  particularly  in  the  margins,  and  the  epithecial  granules 
and  pigment  always  dissolve  completely  in  KOH. 

A  number  of  corticolous  species  belonging  to  the  Lecanora  subfusca 
group  have  been  reported  from  eastern  or  northern  United  States  and,  in 
an  effort  to  show  the  differences  between  these  species  and  L.  chlarotera, 
a  key  to  their  separation  follows. 

This  key  is  mainly  based  on  the  work  of  Magnusson  (1932)  to 
which  I  have  added  information  which  has  been  published  since  then, 
as  well  as  some  of  my  own  observations. 

1.  Apothecial  margin  cortex  50-100  p.  thick,  strongly  gelatinous.  Exclu¬ 
sively  on  the  bark  of  Fagus  (see  Lamb,  1954)  (Nova  Scotia,  Maine) 

. L.  glabrata  Ach. 

1.  Apothecial  margin  cortex  less  developed,  8-35  (-50)  p,  thick . 2 

2.  Cortex  little  developed,  8-15  p.  thick,  KOH—;  crystals  lacking 
in  medulla.  Epithecium  inspersed  with  crystals,  PD  +  red  (see 

Degelius,  1941)  (Maryland,  West  Virginia) . 

. L.  cinereofuscci  Magn. 

2.  Cortex  20-35(-50)  p.  thick,  inner  portion  distinctly  delimited,  KOH 
+  strong  yellow;  medulla  usually  with  heaps  or  clumps  of  crystals 


. 3 

3.  Upper  part  of  hymenium  (epithecium)  coarsely  or  finely  granular.  .  .  5 
3.  Upper  part  of  hymenium  (epithecium)  without  granules,  more  or  less 
reddish  brown . 4 


4.  Apothecia  thick,  urn-like,  margin  coarsely  crenulate  (Maine,  Michi¬ 
gan,  Manitoba) . L.  subrugosa  Nyl. 

4.  Apothecia  thin,  margin  finely  crenulate  or  smooth  (Maine,  New 
York,  Connecticut,  Tennessee,  Michigan,  Oklahoma,  Manitoba, 
Quebec) . L.  subfuscata  Magn. 


5.  On  the  bark  of  conifers . 6 

5.  On  the  bark  of  broadleaf  trees . 7 


6.  Thallus  leprose  or  finely  granular;  epithecium  PD  — ;  disks  dark 
red-brown;  margins  with  a  yellowish  tint  (Maine,  Tennessee)  .... 

. L.  pinastri  (Schaer. )  Magn. 

6.  Thallus  smooth  or  rugose;  epithecium  PD  +  orange  crystals. 

Spores  (13-)  17-20  x  8-13  p. . . 

. L.  insignis  Degel.  (See  discussion  of  L.  degelii.) 

7.  Apothecial  margin  PD  +  red  (substance  unknown)  (Connecticut, 

New  York,  Nova  Scotia,  Saskatchewan,  Quebec) . 

. L.  chlcirona  (Ach.)  Nyl. 

7.  Apothecial  margin  PD  —  or  PD  +  pale  yellow . 8 

8.  Epithecium  PD  +  red,  with  the  production  of  orange  acicular 
crystals  (substance  unknown).  Apothecial  disks  strongly  convex; 
margins  beaded  and  often  discontinuous;  spores  12-14  x  7-8  p.. 

(Long  Island,  North  Carolina,  and  Tennessee) . 

. L.  degelii  Schauer  &  Brodo  (see  p.  217). 


216  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

8.  Epithecium  PD  — .  Apothecial  disks  flat  to  somewhat  convex; 

margins  sometimes  crenulate,  but  always  continuous;  spores  10-13  x 

6-7  jj, . L.  chlarotera  Nyl. 

Lecanora  chlarotera  is  found  on  the  bark  of  various  deciduous  trees 
throughout  the  island. 

Distribution  —  North  American  distribution  largely  unknown,  but 
undoubtedly  common,  at  least  in  the  east;  coastal  Massachusetts,  New 
Jersey,  Arizona;  Europe. 

157.  Lecanora  cinerea  (L.)  Sonim.  Suppl.  FI.  Lapp.  99.  1826.  Lichen 
cinereus  L.  Mantissa  1:132.  1767. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3505B  (10).  SUF¬ 
FOLK  COUNTY:  Brodo  1216  (100A),  2220  (61),  2664  (108),  2745 
(111),  2808  (106),  3355  (62),  3419  (134),  3425  (134),  3433  (134), 
3854  (76);  Orient,  Latham  960,  April  19,  1924  (Latham);  Shelter 
Island,  Latham  24374,  April  1,  1946  (Latham,  MO). 

There  are  two  Long  Island  species  of  Lecanora  in  the  section 
Aspicilia,  L.  cinerea  and  L.  caesiocinerea,  and  they  appear  to  be  very 
closely  related  if,  in  fact,  they  are  not  conspecific. 

In  Europe,  L.  cinerea  and  L.  caesiocinerea  are  separated  easily  by 
their  differing  reactions  with  KOH  and  by  their  pycnoconidia,  which  do 
not  even  come  close  to  overlapping  in  length.  Lecanora  cinerea  has  a 
rapid  KOH  +  yellow  to  red  reaction  (due  to  norstictic  acid)  and  pycno¬ 
conidia  16  to  20;j.  long,  whereas  L.  caesiocinerea  has  a  KOH  —  (or 
dirty  reddish  brown)  reaction  and  pycnoconidia  6  to  12;j.  long  (Hillman 
and  Grummann,  1957). 

On  Long  Island,  the  situation  is  much  more  complicated.  To  begin 
with,  instead  of  two,  there  are  three  divisions  based  on  KOH  reactions. 
KOH  +  red  (norstictic  acid),  KOH  +  yellow  (stictic  acid),  and  KOH — 
Secondly,  there  are  three  size  classes  of  pycnoconidia,  each  class  corre¬ 
lated  with  a  KOH  reaction  type.  Thirdly,  the  three  pycnoconidial  size 
classes  overlap,  especially  in  the  KOH  positive  groups.  The  KOH  +  red 
group  on  Long  Island  has  pycnoconidia  on  the  small  side  of  the  European 
scale,  the  KOH  negative  group  has  pycnoconidia  on  the  large  side  of 
the  scale,  and  the  KOH  +  yellow  group  (the  one  for  which  there  seems 
to  he  no  European  parallel),  introduces  an  intermediate  size  range. 

Since  stictic  acid  and  norstictic  acid  commonly  shift  within  species, 
it  is  reasonable  to  presume  that  the  stictic  acid  specimens  represent  North 
American  chemical  variants  of  the  well  known  L.  cinerea,  which  normally 
produces  norstictic  acid.  The  fact  that  their  pycnoconidial  sizes  overlap 
considerably  adds  to  the  likelihood  of  the  two  variants  being  conspecific. 

The  KOH  —  material  apparently  is  L.  caesiocinerea  with  somewhat 
larger  pycnoconidia  than  seen  in  European  specimens. 

Lecanora  cinerea  is  found  on  exposed  or  partially  shaded  granitic 
rocks  (figure  58). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


217 


Distribution  —  Nova  Scotia,  Maine,  Michigan,  Iowa,  Minnesota, 
Arizona,  Black  Hills,  Washington,  Alaska,  Arctic  Canada;  Arctic-boreal 
element;  Europe;  Asia  ( Zahlbruckner,  1930;  Lynge,  1928). 

158.  Lecanora  conizaea  (Ach.)  Nyl.  Flora  55:  249.  1872.  Lecanora 
expallens  jj.  L.  conizaea  Ach.  Lich.  Univ.  374.  1810. 

Material  seen  —  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 
(1866?)  ( BKL  031909).  NASSAU  COUNTY:  Cold  Spring,  Grout , 
April  1,  1900  (BKL).  SUFFOLK  COUNTY:  Imshaug  25749  (132), 
25770  C  (121  )-Brodo  1706  (133),  2374  (123),  2585  (97),  2831  (115); 
Orient,  Latham  100,  May  23,  1914  (Latham);  Orient,  Latham  22257, 
May  20,  1914  (Latham);  Orient  Point,  Latham,  April  11,  1910  (NYS). 

This  species  is  very  similar  to  Lecanora  symmicta  (Ach.)  Ach.  in 
many  respects.  The  thallus  is  whitish  green  to  yellow-green,  verruculose 
to  granulose,  the  disks  are  yellow  to  buff  or  brown,  and  the  spores  are 
of  the  same  size  and  shape.  In  L.  symmicta,  however,  the  apothecial  mar¬ 
gins  which  are  at  first  smooth,  pale,  and  usually  transluscent,  rapidly 
disappear  with  the  apothecial  disks  becoming  strongly  convex.  Lecanora 
conizaea  has  white  or  thallus-colored,  smooth  apothecial  margins  which 
soon  become  granulose-sorediate,  and  finally  disappear,  leaving  the 
disks  more  or  less  flat  or  slightly  convex.  Some  apothecia  always  show 
the  typical  granulose  lecanorine  margin. 

Lecanora  conizaea  grows  on  the  bark  of  various  trees,  usually  in 
exposed  habitats,  especially  near  the  ocean. 

Distribution  —  Maine,  Tennessee,  North  Carolina,  Black  Hills; 
Europe;  Asia  (Lynge,  1928). 

159.  Lecanora  cupressi  Tuck,  in  Nyl.  Flora  55:  251.  1872. 

Material  seen  —  SUFFOLK  COUNTY:  Montauk,  Latham  3662. 

April  28.  1926  (Latham). 

The  species  was  found  only  once,  and  was  growing  on  wood  of 
what  seems  to  be  Juniperus. 

Distribution  —  Massachusetts  to  Florida  and  Louisiana  (Fink, 
1935):  Temperate  element,  Coastal  Plain  subelement;  endemic. 

160.  Lecanora  degelii  Schauer  &  Brodo,  Nova  Hedw.  11:  528.  1966. 

Material  seen  —  SUFFOLK  COUNTY:  Napeague,  Latham  2847, 

March  1,  1927  (Latham). 

The  Latham  specimen  was  compared  with  the  type  of  L.  insignis 
kindly  sent  to  me  by  Dr.  Degelius,  and  the  two  agreed  in  all  respects 
except  spore  size  and  substrate  type.  The  spores  of  the  Long  Island 
material,  from  oak  bark,  were  smaller  than  those  of  the  type  from  the 
bark  of  Abies. 

Poelt  and  Schauer  discovered  a  correlation  between  spore  size  and 
substrate  in  specimens  of  L.  insignis  collected  recently  by  the  latter  in 
Austria.  Small-spored  specimens  were  from  deciduous  trees  and  larger 
spored  specimens  were  from  coniferous  trees  (Poelt,  and  Schauer,  pers. 
comm.).  Degelius’  specimens  from  the  Smoky  Mountains  showed  the 


218  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

same  correlation.  The  small-spored  population  was  therefore  described  as 
a  new  species  (Schauer  and  Brodo,  1966). 

The  beaded,  almost  disappearing  margin  and  the  production  of 
PD  +  orange  needles  from  the  epithecium  were  both  evident  in  the 
Long  Island  specimen.  The  species  is  very  similar  to  L.  chlarotera,  but 
the  larger  spores  and  PD  +  epithecium  of  L.  degelii  easily  distinguish 
the  two. 

Distribution  —  Smoky  Mountains  of  Tennessee  and  North  Carolina; 
Austria  and  Bavaria  (Schauer  and  Brodo,  1966). 

161.  Lecanora  dispersa  (Pers.)  Somm.  Suppl.  FI.  Lapp.  96.  1826. 
Lichen  dispersus  Pers.  Neue  Ann.  Bot.  1 :  27.  1794. 

Material  seen  —  KINGS  COUNTY:  Brodo  4538  (1).  NASSAU 
COUNTY:  Brodo  3194  (7).  SUFFOLK  COUNTY:  Brodo  2798  (84), 
2839  (115);  Orient,  Long  Beach,  Latham  22332,  December  7,  1944 
( Latham) . 

The  Long  Island  material  of  this  species  agreed  with  both  American 
and  European  specimens  examined  at  the  Farlow  herbarium.  The  apo- 
thecial  margins  of  L.  dispersa  are  usually  described  as  pruinose  or  even 
powdery,  but  in  my  observations  this  is  not  always  the  case. 

The  species  is  similar  to  L.  hageni  in  many  respects,  and  is  often 
included  in  that  species.  Lecanora  hageni,  however,  generally  shows  a 
distinct,  thick  gelatinous  apothecial  margin  cortex,  whereas  L.  dispersa 
is  essentially  without  a  cortex  of  any  kind.  The  latter  seems  to  be 
confined  to  calcareous  rock  and  mortar,  and  the  former  is  most  fre¬ 
quently  found  on  bark.  Both  species  are  commonly  found  growing  with 
species  of  Caloplaca. 

On  Long  Island,  L.  dispersa  was  found  only  on  mortar  and  brick. 
It  has  the  distinction  of  being  the  only  species  found  in  the  western¬ 
most  collection  locality  on  the  island  ...  in  the  heart  of  thickly  populated 
Brooklyn.  It  is  well  known  in  Europe  as  being  a  highly  city-tolerant 
species  (Erichsen,  1957). 

Distribution  —  Michigan,  Indiana,  Minnesota,  Black  Hills,  Arizona, 
Manitoba,  Canadian  archipelago:  Arctic-boreal  element;  circumboreal. 

162.  Lecanora  hageni  (Ach.)  Ach.  Lich.  Univ.  367.  1810.  Lichen 
hageni  Ach.  Lich.  Suec.  Prodr.  57.  1798 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3361  (S  of  97). 

The  similarity  of  this  species  to  L.  dispersa  has  been  discussed  in 
connection  with  the  latter.  Lecanora  hageni  was  collected  only  once,  on 
a  roadside  Ulmus  growing  with  Xanthoria  fallax  and  X.  parietina. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Michigan,  Indi¬ 
ana,  Minnesota,  Black  Hills,  Rocky  Mountains  (seen  in  herb.  MSC), 
Washington,  Alaska:  Temperate  element.  North  Temperate  subelement; 
Europe;  Asia  (Magnusson,  1940). 

163.  Lecanora  laevis  Poelt,  Ber.  Bayer.  Bot.  Ges.  29:  64.  1952. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2829  (115);  Orient, 

Latham  57,  May  23,  1914  (Latham);  Orient,  Latham  817,  October  29, 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


219 


1916  (Latham);  Orient,  Latham  7421,  June  5,  1933  (Latham);  Orient, 
Latham  7424  (22245),  June  5,  1933  (Latham);  Orient,  Long  Beach, 
Latham  3940,  March  27,  1927  (Latham);  Orient,  Latham,  March  3, 
1915  (Latham);  Orient  Point,  Latham  6,  April  18,  1910  (NYS);  Flan¬ 
ders,  S.  Smith  34927,  August  1,  1962  (NYS). 

Lecanora  laevis  bears  an  external  resemblance  to  L.  allophana  (Ach.) 
Nyl.  which,  however,  has  a  thick,  gelatinous  apothecial  margin  cortex 
and  seems  to  be  exclusively  European.  Both  Lamb  (1954)  and  Laundon 
(1958)  have  presented  detailed  descriptions  of  L.  laevis.  On  Long 
Island  it  seems  to  be  restricted  to  the  aerohaline  stratum  on  the  eastern 
Long  Island  coastline. 

Distribution  — -  Nova  Scotia;  southern  Europe  and  North  Africa 
(Lamb,  1954);  Ireland  (Laundon,  1958). 

164.  Lecanora  muralis  (Schreb.)  Rabenh.  Deutschl.  Krypt.  FI.  2:  42. 
1845.  Lichen  muralis  Schreb.  Spic.  FI.  Lips.  130.  1771. 

Material  seen  —  SUFFOLK  COUNTY:  Gardiner’s  Island,  Latham 
36807,  May  23,  1923  (Latham). 

This  species,  common  on  limestone  outcrops  in  the  northeast  and 
elsewhere,  probably  was  introduced  into  Long  Island  with  a  shipment  of 
limestone  building  materials.  It  is  found  nowhere  else  on  the  island, 
probably  due  to  the  lack  of  naturally-occurring  limestone. 

Distribution  —  Connecticut,  central  New  York,  Michigan,  Iowa, 
Minnesota,  Oklahoma,  B'ack  Hills,  Arizona,  Idaho,  Washington,  Alaska: 
Temperate  element.  North  Temperate  subelement;  Europe;  Asia  (Zahl- 
bruckner,  1930;  Magnusson,  1940). 

165.  Lecanora  rubina  (Vill.)  Ach.  Lich.  Univ.  412.  1810.  Lichen 
rubinus  Vill.  Hist.  PI.  Dauph.  3:  977.  1789. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1804  (127),  3443 
(134). 

This  species  was  found  on  exposed  granitic  boulders  within  !4  mile 
of  Long  Island  Sound. 

Distribution  —  Connecticut,  Ontario,  Michigan,  Iowa,  Minnesota, 
Black  Hills,  Arizona,  Idaho,  Washington,  Alaska,  Northern  Saskatche¬ 
wan:  Arctic-boreal  element,  circumboreal  (Ahti,  1964). 

166.  Lecanora  suhintricata  (Nyl.)  Th.  Fr.  Lich.  Scand.  1:  265.  1871. 
Lecanora  varia  var.  suhintricata  Nyl.  Flora  51:  478.  1868. 

Material  seen  —  SUFFOLK  COUNTY:  hnshaug  25616  B  (116): 
Brodo  795  (90B),  2600  (84),  3859  (57). 

Lecanora  suhintricata,  although  listed  by  Hale  and  Culberson 
(1960),  was  not  listed  by  Fink  (1935)  and  has  not  been  mentioned  in 
any  recent  North  American  literature  which  I  have  seen.  The  Long 
Island  material  fits  the  European  descriptions  very  well. 

Lecanora  fuscidula  Degel.  is  a  very  similar  species  from  Maine 
(Degelius,  1940).  I  examined  the  type  specimen  of  L.  fuscidula  kindly 
sent  to  me  by  Dr.  Dege'ius  and  found  it  to  differ  from  L.  suhintricata 
chiefly  in  having  a  well-deve'oped,  gelatinous,  apothecial  margin  cortex, 


220  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

about  16-20[j.  thick.  In  addition,  the  thallus  of  L.  fuscidula  is  scurfy, 
ashy,  and  evanescent,  whereas  that  of  L.  subintricata  is  greenish  and 
granulose,  and  usually  is  well  developed. 

As  in  Europe  (see  Hillman  and  Grummann,  1957)  the  Long  Island 
L.  subintricata  was  found  on  old  wood  and  pine  bark.  It  is  often  asso¬ 
ciated  with  Lecidea  aeruginosa. 

Distribution  — -  Europe;  Asia  (Vainio,  1928). 

167.  Lecanora  symmicta  (Ach.)  Ach.  Syn.  Lich.  340.  1814.  Leca- 
nora  varia  9 .  L.  symmicta  Ach.  Lich.  Univ.  379.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-261  (54). 

Lecanora  symmicta  is  very  similar  to  L.  conizaea.  It  is  placed  with 
Lecanora  rather  than  in  Lecidea,  as  is  often  done,  due  to  its  apparent  close 
ties  with  the  other  members  of  the  Lecanora  varia  group,  many  of  which 
lose  their  margins  in  maturity. 

Lecanora  symmicta  often  resembles  Lecidea  vernalis  in  certain 
respects,  but  the  latter  has  hemispherical  apothecia  in  maturity,  and 
larger  spores  (15-19  fj,  long). 

Distribution  —  Maine,  Connecticut,  North  Carolina,  Tennessee, 
Michigan,  Minnesota,  Black  Hills,  Arizona,  Washington.  Manitoba;  north¬ 
east  Greenland  ( Lynge,  1940c):  Arctic-boreal  element  (?);  Europe; 
Asia  (Vainio,  1928). 

168.  Lecanora  cfr.  varia  (Ehrh.)  Ach.  Lich.  Univ.  377.  1810. 
Lichen  varius  Ehrh.  PI.  Crypt.  Exs.  no.  68.  1785. 

Material  seen  —  SUFFOLK  COUNTY:  Orient,  Long  Beach, 
Latham  3917,  March  27,  1927  (Latham). 

This  specimen  differs  from  all  the  other  Lecanorae  on  Long  Island, 
not  only  in  morphology  but  also  in  substrate  (on  bone).  Bruce  Fink, 
to  whom  this  specimen  was  sent  for  identification  many  years  ago,  called 
it  L.  varia.  The  fact  that  its  apothecial  margin  has  a  well  developed,  gela¬ 
tinous  cortex  puts  it  close  to  L.  varia.  The  Long  Island  specimen,  how¬ 
ever,  contains  atranorin  and  is  therefore  KOH  +  yellow.  Most  authors 
regard  L.  varia  as  a  KOH-  species,  although  some  (e.g.,  Hillmann  and 
Grummann,  1957)  regard  L.  varia  as  either  KOH  +  or  KOH—.  The 
Long  Island  material  also  seems  to  be  very  close  to  Lecanora  sarcopsis 
(Wahlenb.  in  Ach.)  Rohl.  (L.  effusa  fPers.l  Ach.)  which,  however, 
usually  has  an  indistinct,  ungelatinized  apothecial  cortex  and  slightly 
pruinose  apothecial  disks. 

Distribution  —  Connecticut,  North  Carolina,  Indiana,  Iowa,  Wis¬ 
consin,  Minnesota,  Black  Hills,  Arizona,  Washington,  Alaska:  Temperate 
element.  North  Temperate  subelement;  Europe;  Asia  (Lynge,  1928). 

169.  Lecanora  sp. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1189  (101). 

This  specimen  is  in  the  L.  subfusca  group,  and  closely  resembles 
L.  subfuscata  Magn.  except  that  it  does  not  have  large  crystals  in  the 
amphithecium  and  has  a  light  grey  rather  than  a  dark  grey  to  cinereous 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  221 

thallus.  The  thallus  is  granulose  to  almost  sorediate  in  spots  and  smooth 
at  the  edges. 

The  apothecial  disks  are  deep  mahogany  brown,  flat,  0.5-1. Omm  in 
diameter,  with  smooth  to  slightly  crenulate  margins.  The  epithecium  is 
brownish  (remaining  so  in  KOH)  and  is  not  inspersed  with  granules 
(as  in  L.  chlarotera).  The  amphithecium,  although  it  does  not  have 
large,  colorless  crystals,  is  filled  with  smaller  crystals.  The  apothecial 
cortex  is  16-25p,  thick  and  appears  to  be  paraplechtenchymatous.  The 
medulla  and  cortex  are  PD  +  yellow  and  KOH  +  yellow.  The  spores 
are  9-13  x  6-7  tj..  It  was  found  on  the  bark  of  Quercus  alba  near  the 
tree  base. 

43.  OCHROLECHIA  Mass. 

170.  Ochrolechia  parella  (L.)  Mass.  Ricerch.  Auton.  Lich.  32.  1852. 
Lichen  parellus  L.  Mantissa  1:  132.  1767. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25708  (68), 
25853a  (86);  Brodo  59-252  (67),  1112  (78),  1619  (69),  2040  (45), 
2096  (78),  2102  (78),  3809  (66);  Riverhead,  Latham  7787 A,  May  I 
1937  (Latham);  Riverhead,  Latham  36865,  May  16,  1960  (Latham); 
Riverhead,  Latham  36932,  May  16,  1960  (Latham);  Bay  View,  Latham 
36953B,  October  8,  1960  (Latham);  Orient,  Long  Beach,  Latham, 
April  26,  1920  (Latham);  Greenport,  Latham  200,  September  27,  1914 
(FH);  Eastport,  Schrenk  15,  June  24,  1894  (MICH). 

Ochrolechia  parella,  the  commonest  species  of  Ochrolechia  on  Long 
Island,  was  also  found  in  southern  New  Jersey  and  on  Cape  Cod.  It  is 
easily  distinguished  from  the  other  species  of  Ochrolechia  on  Long 
Island  by  its  C  —  thallus  and  its  production  of  variolaric  acid.  This 
substance  is  most  easily  demonstrated  by  introducing  KOH  onto  a 
water  mount  of  apothecial  sections  and  observing  the  thin,  colorless 
needles,  often  in  radiate  clusters,  which  are  produced  in  the  epithecial 
and  amphithecial  regions. 

Verseghy  (1962)  states  that  O.  parella  is  strictly  saxicolous,  but  the 
Long  Island  specimens,  all  corticolous,  compared  favorably  in  mor¬ 
phology  and  chemistry  with  the  type  of  O.  parella  f.  striata  Vers.  (leg. 
Szatala,  Bulgaria,  in  herb.  F).  In  Verseghy’s  monograph,  my  material 
keys  down  to  O.  pallescens  (L.)  Mass,  (see  discussion  under  O.  rosella). 

Ochrolechia  parella  is  found  on  acid  bark  of  both  broad-leaf  and 
coniferous  trees,  most  frequently  in  bogs  (figure  44).  It  has  also  been 
seen  on  old  wood. 

Distribution  —  North  American  distribution  unclear;  Asia  (Zahl- 
bruckner,  1930). 

171.  Ochrolechia  rosella  (Mull.  Arg.)  Vers.  Beih.  Nova  Hedw.  1: 
110.  1962.  Pertusaria  pallescens  var.  rosella  Mull.  Arg.  Flora  62:  483. 
1879. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2147  (102). 

Verseghy  (1962)  published  the  combination  Ochrolechia  rosella, 
using  Pertusaria  pallescens  var.  rosella  Mull.  Arg.  (from  Asia)  as  her 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


222 

basionym.  Tuckerman's  combination,  Lecanora  pallescens  v.  rosella,  was 
listed  as  a  synonym,  but  Verseghy  mistakenly  gave  its  first  date  of  pub¬ 
lication  as  1882  (Syn.  N.  Am.  Lich.  1:  196)  instead  of  1872  (Gen.  Lich. 
125).  Since  Tuckerman’s  combination  precedes  that  of  Muller,  and  the 
epithet  rosella  is  at  the  varietal  level  in  both,  it  would  seem  that  Ver- 
seghy’s  new  combination  should  properly  give  Tuckerman's  name  as  the 
basionym.  Since  it  is  possible  that  Muller’s  rosella  and  Tuckerman’s 
rosella  are  different  species,  especially  in  view  of  the  fact  that  Verseghy 
does  not  list  Tuckerman's  specimens  in  her  notes  on  distribution  and 
apparently  did  not  see  the  Tuckerman  material,  it  is  difficult  to  defend 
considering  Tuckerman’s  name  as  the  true  basionym  of  O.  rosella.  From 
Verseghy’s  description,  except  for  the  single  spore  measurement  given, 
which  is  slightly  large,  it  would  appear  that  the  Long  Island  material  is 
O.  rosella  (Mull.  Arg.)  Vers.;  that  name  will  be  used  until  the  proper 
disposition  of  Tuckerman’s  name  can  be  determined. 

The  Long  Island  specimen  was  identical  (except  for  the  lack  of 
sterile  rays  of  tissue  in  the  apothecia)  with  the  material  filed  under 
Lecanora  pallescens  v.  rosella  in  the  Tuckerman  herbarium.  To  aid  in 
future  discussions  of  the  species,  a  lectotype  should  be  assigned  for 
Tuckerman’s  epithet,  since  he  did  not  cite  any  specimens  in  his  original 
publication,  and  in  1882,  only  wrote  "northern  and  middle  states,  Muhlen¬ 
berg,  etc.”  Fortunately,  Grace  Howard  is  soon  to  publish  a  North  Ameri¬ 
can  monograph  of  Ochrolechia  in  which  Tuckerman’s  var.  rosella  will  be 
typified  (Howard,  pers.  comm.). 

This  species  has  long  been  confused  and  misinterpreted  in  the  litera¬ 
ture.  Although  it  has  generally  been  considered  as  a  variety  of  O.  palles¬ 
cens  (L. )  Mass.,  it  actually  is  not  similar  to  that  species  at  all.  Ochro¬ 
lechia  pallescens  is  apparently  a  relatively  uncommon  oceanic  species  of 
the  British  Isles  and  France  and  has  a  C  —  thallus  and  a  C  — ,  KC  + 
red  apothecial  margin,  with  conspicuously  pruinose  apothecial  disks. 
Ochrolechia  rosella,  according  to  the  material  in  the  Tuckerman  herb¬ 
arium,  has  a  C  +  red  reaction  in  the  thallus  and  apothecial  cortices.  The 
apothecial  disks  are  lightly  or  not  at  all  pruinose,  and  sometimes  show 
distinct  “rays”  of  sterile  tissue  as  described  by  Tuckerman  (1872).  It  is 
a  relatively  common  northeastern  species  and  is  possibly  synonymous 
with  a  very  similar  western  species,  O.  oregonensis  Magn.  These  two 
taxa  are  closely  related  to  O.  tartarea  (L.)  Mass.,  which  also  has  an 
intensely  C  +  red  reaction  in  the  thallus  and  apothecial  cortices.  Accord¬ 
ing  to  Verseghy  (  1962),  O.  tartarea  is  exclusively  saxicolous. 

The  specimen  from  Long  Island  is  very  well  developed.  The  thallus 
is  light  grey  to  whitish,  and  is  rugose  to  thickly  verrucose.  Its  cortex 
is  C  +  red,  but  the  medulla  is  C  —  (and  also  KOH  —  and  PD  — ).  The 
apothecia  are  1 0-20 ( -25 )  mm  in  diameter  and  are  urn  shaped  (i.e.,  with 
a  narrow  stipe  and  basal  attachment).  The  apothecia!  disk  is  yellow- 
orange  to  orange-pink,  lightly  pruinose,  and  appears  very  rough.  No. 
sterile  rays  were  evident.  With  both  C  and  KC.  the  disk  turns  red. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  223 

The  hymenium  is  about  200  p.  thick,  and  the  spores  are  hyaline,  very 
thin  walled,  and  measure  40-60  x  25-26  p. 

The  Long  Island  specimen  was  collected  in  a  bog  on  the  bark  of 
Acer  rubrum. 

Distribution  — -  uncertain:  Temperate  element,  “northern  and  middle 
states”  (Tuckerman,  1882);  Asia  (Verseghy,  1962). 

172.  Ochrolechia  sp. 

Material  seen  —  SUFFOLK  COUNTY :  Greenport,  Latham  793, 
March  28,  1914  (Latham);  Southold,  Latham  973,  March  10,  1922 
( Latham ). 

This  species  has  usually  been  called  O.  pallescens  (L.)  Mass.,  but 
O.  pal'.escens  is  quite  different  in  distribution,  morphology,  and  chem¬ 
istry  (cf.  above). 

In  Verseghy  (1962),  the  material  agrees  fairly  well  with  descrip¬ 
tions  of  both  O.  harmandi  Vers,  and  O.  austroamericana  (Malme)  Vers. 
However,  O.  harmandi  is  known  only  from  Oceania  and  the  Orient,  and 
the  thallus  is  stated  to  be  not  continuous.  Ochrolechia  austroamericana, 
while  agreeing  better  in  thallus  morphology  (continuous,  rugose)  and 
being  more  logical  from  a  phytogeographic  viewpoint  (from  South 
America),  cannot  be  used  for  nomenclatural  reasons.  The  name  is  a  later 
homonym  of  O.  austroamericana  (Ras.)  Ras.  Verseghy  created  the  new 
combination,  apparently  because  the  basionym  of  her  taxon  (O.  tartarea 
var.  austroamericana  Malme,  1937)  has  priority  over  Rasanen's  O.  pal¬ 
lescens  var.  austroamericana  Ras.,  1939) .  Rasanen,  however,  raised  his 
variety  to  species  level  in  1941.  Since  the  Code  of  Botanical  Nomen¬ 
clature  states  that  only  epithets  of  equal  rank  have  priority  over  each 
other,  Rasanen’s  O.  austroamericana  clearly  has  priority  over  Verseghy’s 
combination.  Without  having  examined  any  authentic  material  of  either 
O.  austroamericana  sensu  Verseghy  or  O.  harmandi,  I  decline  from  intro¬ 
ducing  a  new  name,  since  it  may  well  be  that  such  a  common  species 
already  has  a  valid  name. 

This  species,  while  usually  showing  distinctly  pruinose  disks,  often 
lacks  pruina  altogether.  The  C  reaction  of  the  thallus  and  apothecial 
margin  is  confined  to  the  medullary  regions  and  is  negative  in  the 
cortices,  exactly  opposite  from  the  situation  in  O.  rosella. 

It  was  found  on  oak  and  maple  bark. 

Distribution  —  Cape  Cod,  southeastern  United  States,  West  Indies 
(seen  in  herb.  MSC). 

44.  HAEMATOMMA  Mass. 

173.  Haematomma  ochrophaeum  (Tuck.)  Mass.  Atti  I.  R.  Istit. 
Veneto  III.  5:  253.  1860.  Biatora  ochrophaea  Tuck.  Proc.  Amer.  Acad. 
Arts  Sci.  1:  253.  1848. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2125  (102). 

This  species  was  collected  a  number  of  times  in  bogs  on  Cape  Cod 
(Brodo  4174,  4205,  4342,  4371).  Its  morphology,  especially  the  frequent 
lack  of  septation  in  its  spores,  is  discussed  by  Lamb  (1954).  The  species 


224  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

differs  from  Haematomma  sp.  in  ecology  as  well  as  in  morphology,  being 
more  characteristically  found  in  bogs  and  swamps  on  bark  and  wood 
than  in  upland  oak  and  pine  forests. 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  New  Hamp¬ 
shire,  Vermont,  New  York,  North  Carolina,  West  Virginia,  Michigan, 
Ontario,  Quebec:  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit,  Japan  (Culberson,  1963b). 

174.  Cfr.  Haematomma  sp. 

Material  seen  —  SUFFOLK  COUNTY:  42  specimens  collected  by 
Imshaug  and/or  Brodo;  Orient,  Long  Beach,  Latham  22340 ,  December 
7,  1944  (Latham);  Montauk,  Hither  Woods,  Latham  27292,  April  17, 
1946  (Latham). 

The  status  of  this  material  is  far  from  clear.  At  first,  it  appeared 
to  be  identical  with  sterile  material  of  Haematomma  elatinum  (Ach.) 
Mass,  (see  Culberson,  1963b).  With  further  study,  mainly  at  the  sug¬ 
gestion  of  Culberson,  it  became  clear  that  the  soralia  were  entirely  dif¬ 
ferent  (originating  in  irregular  breaks  in  the  thallus,  distinct,  and  puncti- 
form  in  H.  elatinum,  and  in  tiny,  hollow,  globular  to  vermiform  verrucae 
in  this  material),  although  both  can  produce  a  granular  sorediate  crust 
in  the  older  parts  of  the  thallus.  In  addition,  H.  elatinum  is  generally 
found  on  coniferous  bark,  and  H.  sp.  is  found  on  deciduous  bark. 

Pertusaria  trachythallina  also  contains  thamnolic  acid,  and  Imshaug 
(pers.  comm.)  pointed  out  that  several  specimens  of  that  species  show 
vermiform,  hollow,  sorediate  verrucae.  It  seems  odd,  however,  that  no 
smooth,  Pertusaria- like,  sterile  thallus  has  yet  been  found  among  this 
material,  and  that  only  a  few  of  the  dozens  of  fertile  Pertusaria  specimens 
show  any  tendency  towards  the  production  of  hollow  verrucae.  The  dis¬ 
tinctive,  often  thick,  white  to  yellowish,  fibrous  prothalline  margin  seen 
on  almost  every  specimen  of  Haematomma  sp.  contrasts  v/ith  the  absent 
or,  at  most,  very  thin,  white  prothallus  of  Pertusaria  trachythallina. 

Haematomma  leprarioides  (Vain.)  Vain.,  described  from  South 
America,  is  a  similar  species,  usually  found  in  the  sterile  condition.  Its 
soredia  are  farinose,  however,  and  are  produced  in  distinct  punctiform 
soralia  not  associated  with  verrucae. 

With  what  we  now  know  about  this  species,  it  could  as  well  be 
placed  in  Pertusaria  as  in  Haematomma,  and  the  only  reason  for  choos¬ 
ing  the  latter  is  its  superficial  similarity  to  H.  elatinum. 

Distribution  —  Maine,  North  Carolina,  Virginia  (Culberson, 
1963b);  Massachusetts  (Cape  Cod),  New  Jersey:  Temperate  element, 
Appalachian  subelement,  Appalachian  unit  (?);  Europe. 


CANDELARIACEAE 

45.  CANDELARIELLA  Mull.  Arg. 

175.  Candelariella  aurella  (Hoffm.)  Zahlbr.  Cat.  Lich.  Univ.  5: 
790.  1928.  Verrucaria  aurella  Hoffm.  Deutch.  FI.  2:  197.  1796. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  225 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2799  (84),  2840 
(115). 

From  the  descriptions  given  by  Hakulinen  (1954),  these  specimens 
represent  var.  amelia. 

This  species  commonly  grows  in  association  with  species  of  Calo- 
placa  on  mortar  in  exposed  situations. 

Distribution  —  Michigan,  Indiana,  Iowa,  Kansas,  Minnesota,  Black 
Hills,  California,  Washington,  Quebec,  Canadian  archipelago:  Arctic- 
boreal  element  (?);  arctic  and  temperate  Europe  (Hakulinen,  1954); 
Asia  (Magnusson,  1940). 

176.  Candelaria  vitellina  (Ehrh.)  Mull.  Arg.  Bull.  Herb.  Boiss. 
2:  47.  1894.  Lichen  vitellinus  Ehrh.  PI.  Crypt.  Exs.  no.  155.  1785. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1802  (127),  1912 
(114),  2368  (123),  2671  (108),  3441  (134). 

Candelariella  vitellina  was  usually  found  sterile  in  the  Long  Island 
localities,  although  the  few  fertile  specimens  showed  typical  polysporous 
asci.  Most  of  the  Long  Island  specimens  best  fit  the  description  of  var. 
assericola  Ras.  as  given  by  Hakulinen  (1954),  the  thallus  being  granular- 
verruculose,  with  the  granules  or  verrucules  becoming  crowded  into 
flattened  or  rounded  patches  sometimes  becoming  almost  subsquamulose. 
Many  grade  into  what  is  better  referred  to  as  var.  vitellina  with  the 
thalline  granules  and  verrucae  more  dispersed.  For  this  reason,  no  segre¬ 
gation  of  the  Long  Island  material  into  varieties  was  attempted. 

On  Long  Island,  the  species  is  found  on  exposed  granitic  boulders 
associated  with  Sarcogyne  spp.  and  Rinodina  oreina  (figure  68). 

Distribution  —  Maine,  Connecticut,  Michigan,  Indiana,  Wisconsin, 
Minnesota,  Black  Hills,  Arizona,  Washington,  northern  Saskatchewan, 
Manitoba,  Baffin  Island:  Arctic-boreal  element;  circumboreal. 

46.  CANDELARIA  Mass. 

177.  Candelariella  concolor  (Dicks.)  B.  Stein  in  Cohn,  Kryptog.-Fl. 
Schlesien  2(2)  :84.  1879. 

var.  concolor 

Material  seen  —  KINGS  COUNTY:  New  Lots,  Brainerd  (?)  with 
Physcia  millegrana)  (BKL  032039). 
var.  effusa  (Tuck.)  Burnh.  Bryologist  25:  73.  1922. 

Theloschistes  concolor  var.  effuse  Tuck.  Syn.  N.  Amer.  Lich.  1:  52. 
1882. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25581  (52); 
Brodo  59-242  (67),  669  (77),  2424  (118),  2499  (67),  2776  (31),  3146 
(65). 

With  the  exception  of  the  New  Lots  specimen  cited  above  (var. 
concolor),  all  the  Long  Island  material  of  this  species  showed  virtually 
no  foliose  lobes.  It  was  found  growing  on  the  bark  of  various  broad- 
leaf  trees,  usually  at  the  base  or  around  raintracks. 

Distribution  —  Massachusetts,  Connecticut,  central  New  York, 
Arkansas,  Missouri,  Michigan,  Indiana,  Wisconsin,  Minnesota,  Black 


226 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Hills,  Arizona,  Washington:  Temperate  element,  North  Temperate  sub¬ 
element;  Europe;  Asia  (Zahlbruckner,  1930). 

PARMELIACEAE 

47.  PARMELIOPSIS  (Stizenb.)  Nyl. 

178.  Parmeliopsis  aleurites  (Ach.)  Nyl.  Syn.  Lich.  2:54.  1863. 
Lichen  aleurites  Ach.  Lich.  Suec.  Prodr.  117.  1798. 

Material  seen  —  NASSAU  COUNTY:  Brodo  547  (12),  3509 
(10).  SUFFOLK  COUNTY:  37  specimens  collected  by  Imshaug  and/or 
Brodo;  North  Sea,  Latham  36933c,  May  20,  1954  (Latham). 

Parmeliopsis  aleurites  is  found  on  the  bark  of  various  tree  species, 
especially  Pinus  rigida  and  Chamaecy paris  thyoides,  but  also  oaks,  and 
is  occasionally  found  on  lignum.  It  is  most  frequent  in  welldighted  oak 
and  pine  forests. 

Distribution  —  Maine,  Connecticut,  Massachusetts,  New  Jersey, 
North  Carolina  (mountains  and  piedmont),  Tennessee,  Alabama,  Michi¬ 
gan,  Minnesota,  Black  Hills,  Arizona,  boreal  Ontario:  Temperate  element, 
East  Temperate  subelement;  Europe;  Asia  (Vainio,  1928). 

179.  Parmeliopsis  ambigua  (Wulf.  in  Jacq.)  Nyl.  Syn.  Lich.  2:  54. 
1863.  Lichen  ambiguus  Wulf.  in  Jacq.  Coll.  Bot.  4:  239.  1790. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25806  (86), 
25812  (86);  Brodo  1108  (78),  2270  (87). 

Both  usnic  and  divaricatic  acids  were  demonstrated  in  the  Long 
Island  material,  making  it  P.  ambigua  sens.  str.  (or  “chemical  race  A”  in 
the  treatment  by  Culberson  [1955c]). 

Parmeliopsis  ambigua  was  usually  found  on  Chamaecy  paris  (occa¬ 
sionally  on  Pinus)  in  bogs.  As  with  the  other  species  of  Parmeliopsis,  this 
one  seems  to  have  a  strong  specificity  for  conifers  throughout  its  range. 
It  is  found  abundantly  on  pine  on  the  coastal  plain  of  North  Carolina 
(Culberson,  1958a),  on  white  cedars  farther  north,  and  on  spruce  and 
fir  in  boreal  forests. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  New  Jersey, 
Alabama,  Michigan,  Wisconsin,  Black  Hills,  Arizona,  Washington, 
Alaska,  northern  Saskatchewan,  Manitoba,  Baffin  Island,  arctic  Ontario: 
Arctic-boreal  element;  circumboreal. 

180.  Parmeliopsis  placorodia  (Ach.)  Nyl.  Syn.  Lich.  2:55.  1863. 
Parmelia  placorodia  Ach.  Syn.  Lich.  196.  1814. 

Material  seen  —  24  specimens  collected  by  Imshaug  and/or  Brodo; 
Orient,  Latham,  April  1,  1920,  on  Juniperus  (Latham);  Manorville, 
Latham  7767,  May  20,  1937  (Latham);  Riverhead,  Latham  8196,  March 
9,  1938  (Latham);  Napeague,  Latham  8624,  June  1  1,  1938  (Latham); 
Napeague,  Latham  25985,  March  11,  1947  (Latham);  Napeague,  Latham 
34095,  April  1,  1956  (Latham). 

This  species  is  the  most  conspicuous  foliose  member  of  the  pine  bark 
community  (figure  45).  In  some  pine  forests  the  ascending,  often  sub- 
fruticose,  finely-divided  and  abundantly  fruiting  thalli  of  P.  placorodia 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  227 

can  be  seen  on  almost  every  pine  tree,  especially  dead  ones  where  the 
loose  bark  has  ceased  to  slough  off  (pp.  28-29). 

The  substrate  specificity  of  this  species  has  been  discussed  in  detail 
by  Culberson  (1955c).  Parmeliopsis  placorodia  is  almost  exclusively  a 
P/ttMS-dweliing  species,  but  in  various  parts  of  the  country  can  grow  on 
various  species  within  the  genus.  In  the  east,  the  substrate  is  P.  rigida, 
in  the  Great  Lakes  Region  it  is  P.  banksiana,  and  in  the  west  it  is 
P.  ponderosa  (Culberson,  1961b).  On  Long  Island,  Parmeliopsis  placo¬ 
rodia  has  also  been  collected  on  Chamaecy paris  (twice),  Vaccinium 
corymbosum  (once,  cf.  p.  50)  and  on  Qitercus  coccinea-velutina  (twice) 
(not  hybrid;  p.  19).  Rare  occurrences  on  fence  rails  and  shingles  have 
been  noted  as  well. 

Distribution  —  Northeastern  United  States  (map:  Culberson, 
1955c),  Black  Hills,  Arizona,  Ontario:  Temperate  element,  Appalachian 
subelement,  Appalachian-Great  Lakes-Rocky  Mountain  unit;  endemic. 

48.  PARMELIA  Ach. 

181.  Parmelia  appalachensis  W.  Culb.,  Nova  Hedw.  4(3-4):  571. 
1962. 

Material  seen  —  Brodo  59-270  (53). 

This  species  has  long  been  included  in  the  complex  of  pseudocyphel- 
late  Parmeliae  collectively  called  P.  bolliana  Mull .  Arg.  (see  Culberson 
and  Culberson,  1956).  In  his  description  of  the  new  species,  Culberson 
(1962)  indicated  how  it  can  be  separated  from  the  very  similar  P.  fron- 
difera  Merr.  I  have  seen  and  collected  much  material  of  P.  frondifera 
in  central  New  York  (Madison  County,  Bridgeport)  where  it  always  has 
abundant  apothecia  and  an  entirely  pale  undersurface  with  numerous 
pale  buff  rhizines.  The  Long  Island  specimen  has  a  pitch  black  under¬ 
surface  becoming  pale  tan  only  at  the  margins,  and  is  covered  with 
black  or  dark  brown  rhizines.  It  is  essentially  identical  with  an  isotype  of 
P.  appalachensis  (Hale,  Lich.  Amer.  Exs.  63  [MSC]).  The  lobules  so 
characteristic  of  P.  appalachensis  are  not  well  developed  on  the  Long 
Island  specimen  but  are  distinctly  present. 

The  Long  Island  specimen  was  collected  on  the  mossy  base  of  a 
Qitercus  alba  in  an  oak  woods. 

Distribution  —  Nova  Scotia  south  to  North  Carolina  (figure  27): 
Temperate  element,  Appalachian  subelement,  Appalachian  unit  (map: 
Culberson,  1962). 

182.  Parmelia  arseneana  Gyeln.  Ann.  Mycol.  36:  269.  1938. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3025  (50),  3853 

(76),  3870  (62);  Orient,  Latham  942B,  April  25,  1921  (Latham). 

This  species  has  been  variously  treated  by  different  authors.  It 
mainly  comprises  what  Hale  (1955b)  called  Parmelia  conspersa,  chemical 
strain  no.  1. 

It  is  found  on  granitic  boulders. 

Distribution  —  Uncertain. 

183.  Parmelia  aurulenta  Tuck.  Amer.  J.  Sci.  Arts.  II.  25:  424.  1858. 


228  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Material  seen  —  KINGS  COUNTY:  Gowanus,  G.  B.  Brainerd, 
(1866?)  ( BKL  031946  or  031947).  NASSAU  COUNTY:  Brodo  3493 
(4).  SUFFOLK  COUNTY:  Brodo  59-263  (53),  859  (47),  1364  (65), 
1586  (65),  2325  (44),  2495  (67),  3305  (129),  3330  (18),  3916  (54); 
Orient,  Latham,  April  11,  1910  (NYS). 

This  species  is  not  common  on  Long  Island.  It  is  most  frequently 
found  on  the  bark  of  Quercits  alba,  often  at  the  base. 

Distribution  ■ —  Throughout  eastern  United  States  except  for  south¬ 
east  coast  (figure  27):  Temperate  element,  East  Temperate  subelement 
(map:  Hale,  1958);  China,  South  Africa,  India  (Hale,  pers.  comm.). 

184.  Parmelia  caperata  (L.)  Ach.  Meth.  Lich.  216.  1803.  Lichen 
caperatus  L.  Sp.  PI.  1147.  1753. 

Material  seen  —  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 
(BKL  031948);  Cypress  Hill,  Hulst,  1890  (BKL  031949);  Richmond 
Hill,  Hulst,  1890  (BKL  031850).  NASSAU  COUNTY:  Brodo  534  (16). 
SUFFOLK  COUNTY:  96  specimens  collected  by  Imshaug  and/or  Brodo; 
17  specimens  collected  by  Latham  (Latham). 

Parmelia  caperata,  one  of  the  most  common  species  on  Long  Island, 
was  used  in  various  ecological  and  experimental  studies. 

Often,  soredia  are  scarcely  produced  at  all  or  are  in  minute,  almost 
isidiate  heaps  of  granules  scattered  over  the  thallus  surface.  The  lobes 
are  almost  always  broad  and  rounded,  but  on  rare  occasions  they 
become  laciniate. 

Parmelia  caperata  is  most  common  on  tree  bases  of  almost  any 
species  of  tree  on  Long  Island.  It  is  also  found  with  fair  frequency  on 
exposed,  partially  shaded,  or  shaded  boulders,  even  in  the  salt  spray 
zone  near  the  coast  (p.  60). 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  New  Jersey, 
North  Carolina,  Tennessee,  Alabama,  Arkansas,  Missouri,  Oklahoma, 
Indiana,  Michigan,  Ontario,  Wisconsin.  Minnesota,  Black  Hills,  Arizona, 
Washington,  Manitoba:  Temperate  element.  North  Temperate  subele¬ 
ment;  Europe;  Asia  (Asahina,  1952). 

185.  Parmelia  conspersa  (Ach.)  Ach.  Meth.  Lich.  205.  1803. 
Lichen  conspersus  Ach.  Lich.  Suec.  Prodr.  118.  1789. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25602  (SW.  of 
106);  Brodo  1908  (114),  2379  (123),  2655  (108),  2709  (111),  3018 
(17),  3024  (50),  3029  (50),  3381  (94),  3421  (134),  3444  (134), 
3846  (76);  Orient,  Latham  942 A,  April  25,  1921  (Latham);  Shelter 
Island,  Latham  22929,  October  14,  1949  (Latham). 

This  species  is  the  isidiate  member  of  the  Xanthoparmeliae  which  was 
considered  under  the  name  P.  isidiata  (Anzi)  Gyeln.  by  Hale  (1955b, 
1956a)  and  later  (Hale,  1961a)  as  P.  lusitana  Nyl.  Hale  (1964)  dis¬ 
cusses  the  systematics  of  this  and  related  taxa  in  great  detail.  It  is  dis¬ 
tinct  from  P.  plittii  Gyeln.  in  the  color  of  its  undersurface:  black  to 
very  dark  brown  in  P.  conspersa  and  pale  brown  to  buff  in  P.  plittii.  All 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  229 

of  the  Long  Island  material  of  both  species  contained  both  norstictic  and 
stictic  acids. 

Parmelia  conspersa  is  found  on  exposed  granitic  boulders,  as  are 
most  of  the  other  Xanthoparmeliae.  Several  members  of  this  group  are 
often  found  growing  together,  even  intermingling  thalli,  and  one  must 
be  very  careful  in  order  to  get  an  unmixed  collection. 

Distribution  —  Eastern  North  America,  southern  Canada,  Black 
Hills,  Oregon,  California  (map:  Hale,  1964):  Temperate  element, 
North  Temperate  subelement;  Europe. 

186.  Parmelia  dilatata  Vain.  Acta  Soc.  Faun.  FI.  Fenn.  7(7):  33. 
1890. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1760  (127),  1869 
(117). 

This  species  is  distinctive  since  few  other  marginally  sorediate 
Parineliae  contain  protocetraric  acid.  It  generally  has  been  mentioned 
in  the  literature  under  the  name  Parmelia  robusta  Degel.  Hale  (1965b) 
discusses  its  nomenclatural  problems. 

Parmelia  dilatata  is  usually  a  very  broad,  vigorous  plant.  Both  Long 
Island  specimens,  however,  were  small  lobed  (3-4  mm  broad).  They 
were  found  on  Quercus  velutina  in  the  fog  belt  region  of  the  island’s 
south  fluke. 

Distribution  —  South  Carolina,  Georgia,  Florida,  tropic  and  sub¬ 
tropic  regions  (Hale,  1959a):  Tropical  element.  Coastal  Plain  subelement; 
Europe,  Asia  (Hale,  1965b). 

187.  Parmelia  galbina  Ach.  Syn.  Meth.  Lich.  195.  1814. 

Material  seen  —  KINGS  COUNTY:  Gowanus,  G.  B.  Brainerd, 

(1866?)  (BKL).  NASSAU  COUNTY:  Cold  Spring,  Grout,  April  1, 
1900  (BKL).  SUFFOLK  COUNTY:  22  specimens  collected  by  Imshaug 
and/or  Brodo;  Orient,  Latham  3,  May  2,  1914  (Latham). 

Culberson  (1961c)  in  his  study  of  the  Parmelia  quercina  group, 
presented  an  excellent  description  and  discussion  of  this  species  including 
a  photograph  of  the  characteristic  nioniliform  cells  of  the  medulla. 

The  species  is  clearly  a  member  of  the  breast  height  community  on 
Quercus  velutina  (figure  51). 

Distribution  —  Temperate  element.  East  Temperate  subelement, 
Japan  (map:  Culberson,  1961c). 

188.  Parmelia  hypotropa  Nyl.  Syn.  Lich.  1:  378.  1860. 

Material  seen  —  SUFFOLK  COUNTY:  26  specimens  collected  by 

Imshaug  and/or  Brodo;  15  specimens  collected  by  Latham  (Latham); 
Orient  Point,  Latham ,  October  11,  1909  (NYS);  Orient  Point,  Latham, 
April  18,  1910  (NYS);  Orient  Point,  Latham,  April  25,  1910  (NYS); 
Orient  Point,  Latham  15,  April  4,  1910,  April  11,  1910,  April  4  and  18, 
1910  (Note:  three  packets)  (NYS,  MICH);  Orient,  Latham  3926, 
March  27,  1927  (NYS?). 

Parmelia  hypotropa  can  be  confused  with  several  closely  related 
Amphigymniae,  especially  P.  perlata  and  P.  perforata,  or  even  with 


230 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


P.  reticulata  (subgenus  Hypotrachyna) .  The  table  below  summarizes  the 
distinctions  between  these  species. 


P.  hypotropa 

P.  perforata 

P.  perlata 

P.  reticulata 

1.  soredia 

marginal 

absent 

submarginal 

marginal  and 

sometimes 

submarginal 

2.  irregular 
white  margin 
on  under- 

usually 
conspicuous; 
rarely  scanty 

as  in  P. 
hypotropa 

absent 

absent 

surface 

3.  revolute 

absent 

absent 

present 

absent 

margins 

4.  undersurface 

smooth 

smooth 

minutely 

rugulose 

smooth 

5.  uppersurface 

occasionally 
with  scattered 
white  maculae 

uniform  or 
occasionally 
with  scattered 
white  maculae 

uniform 

with 

conspicuous 
reticulate 
cracks  or 
maculae,  esp. 
on  older 
portions 

6.  Chemistry, 

norstictic  and 

norstictic 

stictic 

salacinic 

other  than 

stictic  acids 

alone 

alone 

acid  alone 

atranorin 

7.  cilia 

>  1  mm; 

>  1  mm; 

<  1  mm; 

<  1  mm; 

conspicuous 

conspicuous 

inconspicuous 

inconspicuous 

8.  apothecia 

not  seen 

common; 

not  seen 

not  seen 

( rare ) 

perforate 

( rare ) 

( rare ) 

9.  abundance 

common 

common 

very  rare 

infrequent 

Parmelia  hypotropa  is  a  photophilous  species,  being  most  commonly 
collected  on  the  bark  or  twigs  of  various  trees  in  exposed  areas,  especially 
where  the  humidity  is  maintained  at  a  fairly  high  level.  It  is  best  developed 
on  trees  in  the  oceanic  dune  areas  of  the  island’s  south  fluke,  especially 
in  hollows  and  on  the  lee  sides  of  the  dunes,  but  frequently  occurs  on 
exposed  bog  trees  as  well  (figure  41).  It  occasionally  is  found  in  oak 
forests. 

Distribution  —  Mainly  Ozark  and  southern  Appalachian  Mountains 
(Hale,  1961a),  but  also  California  and  Mexico  (Hale,  1965b):  Tem¬ 
perate  element,  Appalachian  subelement,  Appalachian-Ozark  unit;  Eu¬ 
rope,  Africa,  Asia  (Hale,  1965b). 

189.  Parmelia  livida  Tayl.  Lond.  J.  Bot.  6:  171.  1847. 

Material  seen  —  NASSAU  COUNTY:  Brodo  556  ( 13),  558  ( 13) ; 
Massapequa,  S.  Cain  39,  40,  June  20,  1935  (NY).  SUFFOLK  COUNTY: 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


231 


39  specimens  collected  by  Imshaug  and/or  Brodo;  Northwest,  Latham 
26136C,  April  17,  1947  (Latham);  Northwest,  Latham  26136,  April  10, 
1947  (MO).  (Note:  specimen  numbers  and  dates  are  as  on  the  original 
labels  except  for  the  segregate  designation  “C”  in  the  preceding  specimen.) 

This  species  has  been  discussed  in  detail  by  Culberson  (1961c).  It 
is  outwardly  very  similar  to  P.  galbina,  from  which  it  can  be  separated 
by  its  uniformly  white  medulla,  PD  —  and  KOH  +  red-brown  reactions, 
and  its  lack  of  medullary  moniliform  cells.  Hale  (pers.  comm.)  also  draws 
attention  to  the  difference  in  the  rhizines  of  the  two  species:  branched  in 
P.  livida  and  simple  in  P.  galbina. 

Parmelia  livida  grows  on  the  bark  of  various  species  of  trees,  usually 
at  breast  height,  in  oak  forests. 

Distribution  —  Throughout  southeastern  United  States,  northward 
along  the  east  coast  to  New  Hampshire  (map:  Culberson,  1961c):  Tem¬ 
perate  element,  East  Temperate  subelement;  endemic. 

190.  Parmelia  michauxiana  Zahlbr.  Cat.  Lich.  Univ.  6:  244.  1929. 
Parmelia  epiclada  Hale,  Bryologist  62:  125.  1959. 

var.  michauxiana 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-216  (68),  620 
(39),  1191  (  101),  1421  (83),  1522  (100B),  1784  (127),  1883  (117), 
1901b  (114),  2222  (61),  3249  (119),  3256  (119),  3262  (119),  3907 
(112);  Orient,  Latham,  April  25,  1921  (Latham);  Napeague,  Latham 
8121,  November  6,  1938  (Latham);  Northwest,  Latham  26135B,  April 
10,  1947  (Latham), 
var.  laciniata  (Hale)  comb.  nov. 

Parmelia  epiclada  var.  laciniata  Hale,  Bryologist,  62:  126.  1959. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1191  (101). 

Although  this  species  is  considered  to  be  a  member  of  the  subgenus 
Hypotrachyna,  it  has  many  characteristics  in  common  with  certain 
species  in  Amphigymnia.  For  example,  its  lobes  are  often  very  broad 
(up  to  5  mm  across)  and  bear  sparse  but  distinct  black  marginal  cilia. 
On  rare  occasions,  specimens  will  be  encountered  having  very  narrow 
lobes  (mostly  1-2.5  mm  broad)  curled  inward  and  ascending.  These 
specimens  can  be  called  var.  laciniata.  Apothecia  are  commonly  present 
but  are  never  perforate.  Protocetraric  acid  and  atranorin  are  always  pres¬ 
ent  and  the  medulla  is  conspicously  thick  and  very  cottony.  This  com¬ 
bination  of  characters  is  usually  sufficient  to  separate  it  from  any  similar 
species  on  Long  Island. 

Parmelia  michauxiana  is  a  member  of  the  breast  height  community 
on  oak. 

Distribution  —  Temperate  element.  Coastal  Plain  subelement  (see 
Ha'e,  1 959b) ;  endemic. 

191.  Parmelia  olivetorum  Nyl.  Not.  Sallsk.  Faun.  FI.  Fenn.  For- 
handl.,  n.  ser.  8:  180.  1866. 


232  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Material  seen  —  SUFFOLK  COUNTY :  Fisher’s  Island,  Latham, 
June  24,  1929,  (Latham);  Montauk,  Latham  36782,  July  5,  1931 
(Latham) . 

This  species  must  be  extremely  rare  on  the  island,  since,  although 
1  made  a  special  effort  to  find  specimens  in  the  two  localities  listed 
above,  1  never  saw  a  trace. 

Among  the  taxonomic  problems  involved  in  this  species  is  the 
controversy  concerning  the  logic  in  recognizing  species  solely  on  the 
basis  of  chemical  differences.  Parmelia  olivetorum  contains  atranorin 
and  olivetoric  acid,  and  closely  related  P.  cetrarioides  contains  atranorin 
and  perlatolic  acid  (Culberson,  1962).  Culberson  (1958b)  discussed 
these  chemical  populations  in  detail,  presenting  maps  of  their  distribu¬ 
tion.  The  two  populations  both  have  Appalachian-Great  Lakes  distribu¬ 
tions,  although  P.  olivetorum  seems  to  have  more  northern  tendencies 
(Culberson,  1958b).  Nomenclaturally,  whether  one  considers  the  two  as 
synonymous  or  as  distinct  species,  the  name  which  must  be  used  for 
the  Long  Island  material  (which  contains  olivetoric  acid)  is  P.  olivetorum. 

Latham’s  specimens  were  found  on  rock. 

Distribution  —  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit  (map:  Culberson,  1958b);  Europe;  Asia  (ibid). 

192.  Parmelia  perforata  (Wulf.  in  Jacq.)  Ach.  Meth.  Lich.  217. 
1803.  Lichen  perforatus  Wulf.  in  Jacq.  Coll.  Bot.  1:  116,  pi.  3.  1786. 

Material  seen  —  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 
(1866?)  ( BKL  031952);  Ridgewood,  G.  B.  Brainerd,  (BKL).  NASSAU 
COUNTY:  Cold  Spring,  Grout,  April  1,  1900  (BKL);  Cold  Spring, 
Harris,  April  28,  1904  (MICH).  SUFFOLK  COUNTY:  35  specimens 
collected  by  Imshaug  and/or  Brodo;  16  specimens  collected  by  Latham 
(Latham);  Eastport,  Schrenk,  June  24,  1894  (MICH);  Sayville,  Lloyd, 
135,  December  2,  1896  (NY);  Yaphank,  Davis,  June  3,  1929  (STATEN 
ISLAND);  Flanders,  Latham  7232,  February  3,  1933  (MICH). 

All  the  specimens  collected  on  Long  Island  showed  the  presence  of 
norstictic  acid.  Until  recently,  it  was  believed  that  P.  perforata  was  char¬ 
acterized  by  containing  salacinic  acid  and  could  be  separated  on  this 
basis  from  P.  hypotropoides  Will.,  which  contains  norstictic  acid  (Hale, 
1957c).  However,  Hale  (pers.  comm.)  more  recently  concluded  that 
specimens  earlier  determined  as  P.  perforata  are  P.  cetrata  Ach.,  and  that 
P.  hypotropoides  is  synonymous  with  P.  perforata.  Parmelia  erecta  Berry 
is  also  a  synonym  of  P.  perforata. 

Parmelia  perforata  is  remarkably  similar  to  P.  hypotropa  in  many 
respects,  and  the  two  are  undoubtedly  closely  related  (p.  229). 

Parmelia  perforata  occurs  mainly  in  pine-oak  forests  and  bogs,  and 
like  P.  hypotropa,  is  found  on  exposed  trees  and  shrubs  in  the  humid 
oceanic  habitats  in  eastern  Long  Island  (figure  52). 

Distribution  —  Temperate  element.  East  Temperate  subelement 
(map:  Hale,  1957c);  Ireland,  Madagascar  (Hale,  1965b). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  233 

193.  Pctrmelia  perlata  (Huds.)  Ach.  Meth.  Lich.  216.  1803.  Lichen 
perlatus  Huds.  FI.  Angl.  448.  1762. 

Material  seen  —  SUFFOLK  COUNTY :  Orient  Point,  Latham, 
April  18,  1910  (NYS);  Orient  Point,  Latham,  April  25,  1910  (Note: 
mixed  with  P.  hypotropa)  (NYS). 

An  excellent  description  of  this  species  and  a  discussion  of  its  nomen¬ 
clature  has  been  provided  by  Hale  (1961b).  Parmelia  perlata  is  com¬ 
pared  with  more  common,  similar  species  in  the  discussion  of  P.  hypo¬ 
tropa.  Its  occurrence  on  Long  Island  extends  the  known  range  of 
P.  perlata  northward  from  the  southern  Appalachians. 

Distribution  —  Unglaciated  southern  Appalachians  (Hale,  1961b): 
Tropical  element,  Appalachian-Temperate  subelement;  South  America, 
Mexico,  Japan,  and  Australia  (ibid). 

194.  Parmelia  plittii  Gyeln.  Fedde.  Repert.  29:  287/415.  1931. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  878  (47),  1805 

(127),  3879  (62);  Shelter  Island,  Latham  22929A,  October  14,  1949 
(Latham). 

Parmelia  plittii  is  separated  from  P.  conspersa  mainly  by  the  color 
of  the  thallus  undersurface  (see  under  P.  conspersa) .  Hale  (1964)  also 
points  out  that  P.  plittii  never  shows  the  loosely-attached  form  which  is 
often  found  in  P.  conspersa.  The  ecology  as  well  as  the  distribution  of  the 
two  species  on  Long  Island  appear  to  be  identical. 

Distribution  —  Widespread  in  tropical  America,  and  in  Africa, 
Appalachians-Great  Lakes  region  (map:  Hale,  1964):  Tropical  element, 
Appalachian-Temperate  subelement. 

195.  Parmelia  reticulata  Tayl.  in  Mack.  FI.  Hibern.  2:  148.  1836. 

Material  seen  —  KINGS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 

(1866?)  ( BKL  031954).  SUFFOLK  COUNTY:  Brodo  1765  A  (127), 
2136  (102),  3264  (119). 

The  reticulate  cracks  which  seem  to  first  appear  as  white  reticulate 
maculae  are  not  always  very  conspicuous.  They  are  usually  best  developed 
on  the  older  portions  of  the  thallus.  The  black  rhizines  are  often  long  at 
the  margins  and  extend  out  from  under  the  thallus,  giving  the  appear¬ 
ance  of  marginal  cilia.  Under  these  conditions,  P.  reticulata  bears  several 
resemblances  to  P.  hypotropa,  and  their  separation  is  summarized  under 
the  latter  species. 

Of  the  three  Long  Island  collections,  two  were  on  Quercus  velutina 
in  oak  woods  and  one  was  on  Acer  rubrum  in  a  white  cedar  bog.  All 
three  specimens,  however,  were  found  in  the  humid  and  oceanic  south 
fluke  region  of  the  island. 

Distribution  —  North  Carolina,  Tennessee,  Alabama,  Arkansas, 
Missouri,  Oklahoma,  Arizona;  Great  Lakes  region  (seen  in  herb.  MICH); 
western  United  States  (Hale,  pers.  comm.);  “cosmopolitan”  (Hale, 
1961a):  Temperate  element  (?),  North  Temperate  subelement  (?); 
Europe,  Asia,  Africa,  Australia  ( Zahlbruckner,  1930). 


234  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

196.  Parmelia  rudecta  Ach.  Syn.  Meth.  Lich.  197.  1814. 

Material  seen  —  SUFFOLK  COUNTY :  84  specimens  collected  by 

Imshaug  and/or  Brodo;  14  specimens  collected  by  Latham  (Latham); 
Orient,  Young  (BKL);  Orient  Point,  Latham,  April  11,  1910  (NYS); 
Southold,  (Davis?) ,  Sept.  5,  1912  (STATEN  ISLAND). 

This  very  common  species  shows  a  great  deal  of  variation  in  the 
extent  of  isidial  production.  A  few  specimens  were  almost  completely 
devoid  of  isidia,  but  the  great  majority  showed  the  typical  coralloid  form 
or  the  somewhat  flattened  type  described  by  Culberson  (1962). 

The  ecological  limits  of  P.  rudecta  are  broad.  It  is  found  on  a 
variety  of  phorophyte  species  and  commonly  grows  on  the  base  as  well 
as  at  breast  height.  It  is  abundant  throughout  the  island  except  for  its 
sharply  delimited  distribution  at  the  Nassau-Suffolk  County  border  which 
presumably  is  due  to  the  city-effect. 

Distribution  —  Throughout  eastern  United  States  (map:  Culberson 
&  Culberson,  1956):  Temperate  element.  East  Temperate  element;  China, 
Argentina  (Culberson,  1962);  Japan,  Africa  (Hale,  1965a). 

197.  Parmelia  saxatilis  (L.)  Ach.  Meth.  Lich.  204.  1803.  Lichen 
saxatilis  L.  Sp.  PI.  1  142.  1753. 

Material  seen  —  COUNTY  UNKNOWN:  Long  Island,  Austin 
(BKL  031946).  NASSAU  COUNTY:  Brodo  533  (16),  543  (12),  565 
(11),  1512  (14).  SUFFOLK  COUNTY:  81  specimens  collected  by 
Imshaug  and/or  Brodo;  10  specimens  collected  by  Latham  (Latham). 

Due  to  its  abundance,  P.  saxatilis  was  used  in  a  number  of  eco¬ 
logical  studies  (Brodo,  1961a).  The  species  has  a  significant  specificity 
for  Quercus  velutina  (including  Q.  coccinea:  see  p.  21)  in  pine-oak 
and  in  scarlet-black  oak  forests.  It  is  most  conspicuous  in  the  latter  vege¬ 
tation  type,  as  is  the  very  closely  related  P.  sulcata  (table  9).  Although 
the  two  species  are  almost  always  found  in  the  same  oak  stand  they 
are  often  present  in  widely  different  quantities.  In  two  typical  oak  stands 
in  central  Long  Island,  P.  sulcata  outnumbered  the  thalli  of  P.  saxatilis 
by  a  large  margin  in  one  stand,  and  was  essentially  absent  from  another 
stand  in  which  P.  saxatilis  was  very  abundant.  The  principle  of  non-over- 
lapping  niches  of  closely  related  species  may  play  a  part  in  this  curious 
distribution  (Brodo,  1961a). 

Parmelia  saxatilis,  besides  being  found  on  various  trees,  also  grows 
on  boulders  and  on  rare  occasions,  even  on  soil. 

Distribution  —  Throughout  temperate,  arctic,  and  boreal  North 
America,  including  northern  Saskatchewan,  Manitoba,  arctic  Ontario, 
Canadian  east  arctic,  and  Baffin  Island:  Arctic-boreal  element:  circum- 
boreal. 

198.  Parmelia  stenophylla  (Ach.)  Heug.  Correspondzbl.  Naturf. 
Verein.  Riga  8:  109.  1855.  Parmelia  conspersa  (3.  P.  stenophylla  Ach. 
Meth.  Lich.  206.  1803. 

Material  seen  —  KINGS  COUNTY:  Gowanus,  G.  B.  Brainerd 
(1866?)  (BKL  031951).  SUFFOLK  COUNTY:  Imshaug  25691  (72); 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  235 

Brodo  2667  (108);  Northwest,  Latham,  April  10,  1947  (Latham);  Shel¬ 
ter  Island,  Latham  24375,  April  1,  1941  (Latham). 

The  circumscription  of  Parmelia  stenophylla  is  still  not  clear  (see 
discussion  under  P.  tasmanica) .  The  Long  Island  specimens  having  no 
isidia  and  a  pale  lower  surface  all  contain  salacinic  acid  and  are  more  or 
less  loosely  attached.  They  represent  the  most  “typical”  of  the  P.  steno¬ 
phylla  populations. 

Parmelia  stenophylla  is  found  on  exposed  or  partially  shaded  granitic 
boulders  throughout  the  morainal  regions  of  the  island. 

Distribution  —  Throughout  United  States,  southern  Canada,  and 
a  few  arctic  localities  (map:  Hale,  1955b) :  Temperate  element  (?),  North 
Temperate  subelement;  Europe;  Asia  (Magnusson,  1940). 

199.  Parmelia  subaurifera  Nyl.  Flora  66:  22.  1873. 

Material  seen  —  SUFFOLK  COUNTY :  65  specimens  collected 
by  Imshaug  and/or  Brodo;  Greenport,  Latham,  January  26,  1923  (La¬ 
tham);  East  Marion,  Latham  27,  May  3,  1914  (Latham);  Orient,  Latham 
3928,  March  27,  1927  (Latham);  Orient,  Latham  7454,  June  5,  1933 
(Latham);  Orient,  Latham  8584,  April  30,  1939  (Latham);  Northwest, 
Latham  27214,  April  17,  1947  (Latham);  Riverhead,  Latham  36889, 
May  25,  1960  (Latham). 

Parmelia  subaurifera  is  the  only  Melanoparmelia  on  Long  Island. 
It  grows  on  various  types  of  tree  bark  in  a  variety  of  vegetation 
types.  One  specimen  was  found  growing  on  an  exposed  boulder  in  the 
Montauk  area. 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  Connecticut, 
New  Jersey,  Tennessee,  Michigan,  Ontario,  Wisconsin:  Temperate  ele¬ 
ment,  Appalachian  subelement,  Appalachian-Great  Lakes  unit;  Europe, 
Asia  (Vainio,  1928). 

200.  Parmelia  subrudecta  Nyl.  Flora  69:  320.  1886. 

Material  seen  —  NASSAU  COUNTY:  Brodo  561  (13),  1496  (9). 
SUFFOLK  COUNTY:  Brodo  868  (47),  1288  (21),  1292  (19),  1440 
(83),  2197  (20),  2350  (42),  2478  (23),  3039  (50),  3109  (68), 
3221  (35),  3237  (35). 

This  species,  long  considered  under  the  name  P.  borreri  (Turn.  ex.  Sm. 
in  Sm.  &  Sowerby)  Turn,  or  P.  dubia  (Wulf.  in  Jacq.)  Schaer.  (see  Culber¬ 
son,  1962  and  Hale,  1959b),  is  properly  P.  subrudecta.  Parmelia  borreri 
has  a  black  undersurface  and  contains  gyrophoric  acid,  whereas  P.  sub¬ 
rudecta  has  a  pale  undersurface  and  contains  lecanoric  acid  (Hale, 
1965a). 

Parmelia  subrudecta  seems  to  have  a  strong  affinity  for  the  mature 
red  oak  forests  of  the  morainal  regions  (figure  63).  It  is  found  on  the 
bark  of  various  deciduous  trees  in  shaded  woods. 

Distribution  —  Throughout  the  Appalachian  mountains  and  the 
Great  Lakes  region,  Arkansas,  New  Mexico,  Colorado,  California,  Mex¬ 
ico  (map:  Hale,  1965a);  Arizona:  Temperate  element,  Appalachian  sub- 


236  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

element,  Appalachian-Great  Lakes-Rocky  Mountains  unit;  Europe,  South 
Africa,  and  Australia  (Hale,  1965a);  eastern  Asia  (Culberson,  1962). 

201.  Parmelia  sulcata  Tayl.  in  Mack.  FI.  Hibern.  2:  145.  1836. 

Material  seen  —  QUEENS  COUNTY;  Ridgewood,  G.  B.  Brain- 

erd  (?)  ( BKL  031956).  NASSAU  COUNTY:  Brodo  535  (16),  1315 
(15).  SUFFOLK  COUNTY:  83  specimens  collected  by  Imshaug  and/or 
Brodo;  Riverhead,  Latham,  May  1,  1960  (Latham);  Riverhead,  Latham, 
May  16,  1960  (Latham);  Greenport,  Latham,  May  12,  1960  (Latham); 
Orient,  Latham  696,  March  30,  1914  (Latham);  Orient,  Latham  8583, 
April  30,  1939  (Latham);  Northwest,  Latham  26136D,  April  17,  1947 
(Latham);  Orient  Point,  Latham,  April  18,  1910  (NYS). 

The  ecology  of  Parmelia  sulcata  has  been  discussed  with  P.  saxatilis, 
which  it  closely  resembles  both  morphologically  and  ecologically.  Par¬ 
melia  sulcata  also  shows  a  significant  association  with  Quercus  velutina, 
especially  in  the  pine-oak  forests  (Brodo,  1961a). 

Distribution  —  Nova  Scotia,  Maine,  Masaschusetts,  Connecticut, 
New  Jersey,  North  Carolina,  Michigan,  Ontario,  Wisconsin,  Minnesota, 
Black  Hills,  Arizona,  Washington,  Alaska,  British  Columbia,  Saskatche¬ 
wan,  Manitoba,  Quebec,  Baffin  Island:  Arctic-boreal  element;  circum- 
boreal;  listed  as  having  an  Appalachian-Great  Lakes-Rocky  Mountain 
distribution  by  Hale  (1961a). 

202.  Parmelia  tasmanica  Hook,  and  Tayl.  Lond.  J.  Bot.  3:  644. 
1844. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1906  (114),  2369 
(123),  3080  (128);  Southold,  Janning’s  Woods,  Latham  63868  (36868), 
July  4,  1933  (Latham). 

Members  of  the  Parmelia  stenophylla  group  having  a  black  lower 
surface  and  salacinic  acid  can  be  referred  to  this  species.  It  was  previously 
designated,  at  least  in  part,  as  P.  conspersa,  chemical  strain  no.  2,  by 
Hale  (  1955b).  Degree  of  adnation,  formerly  considered  by  Hale  (1955b, 
1956a)  to  be  an  important  differentiating  criterion,  appears  to  be  un¬ 
reliable.  Loosely  attached  specimens,  which  by  former  standards  would 
have  all  been  called  P.  stenophylla  and  expected  to  show  a  pale  lower 
surface,  now  are  found  to  have  black  lower  surfaces  in  some  specimens  and 
pale  lower  surfaces  in  others.  Abandoning  degree  of  adnation  as  a  prime 
character,  the  taxa  can  he  separated  on  the  basis  of  lower  surface  color 
alone,  and  this  has  been  done  with  many  of  the  specimens  annotated 
by  Hale  in  the  Michigan  State  University  Herbarium.  Whether  or  not 
these  taxa  should  be  recognized  at  the  species  level  is  a  matter  for  future 
discussion  and  investigation. 

The  species  is  strictly  saxicolous,  usually  on  exposed  boulders. 

Distribution  —  Eastern  United  States,  Japan,  Australia,  Europe 
(Hale,  pers.  comm.). 

49.  HYPOGYMNIA  Nyl. 

By  virtue  of  their  hollow  thalli,  complete  lack  of  rhizines,  and 
distinct  chemistry  (Krog,  1951),  members  of  the  well-defined  su'ogenus 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  237 

Hypogymnia  of  the  genus  Parmelia  seem  to  be  sufficiently  distinct  to  be 
considered  together  as  a  separate  genus. 

203.  Hypogymnia  physodes  (L.)  Nyl.  Lich.  Paris  39.  1896.  Lichen 
physodes  L.  Sp.  PI.  1144.  1753. 

Material  seen  —  SUFFOLK  COUNTY :  46  specimens  collected  by 
Imshaug  and/or  Brodo;  10  specimens  collected  by  Latham  (Latham). 

Hypogymnia  physodes  is  a  common  species  on  Long  Island  and,  as 
with  most  common  species,  shows  a  great  deal  of  morphological  varia¬ 
tion.  The  lobes  can  be  long  and  slender,  or  rather  short,  broad,  and  fan 
shaped.  The  soredia  occur  in  abundant  labriform  soralia  bursting  from 
the  tips  of  hollow  lobes,  or  soredia  are  almost  entirely  absent.  It  is  inter¬ 
esting  to  note  that  when  the  lobes  are  narrow,  soralia  appear  to  be  abun¬ 
dant,  whereas  in  broad-lobed  forms,  the  soralia  are  very  scanty. 

Hypogymnia  physodes  was  found  on  the  bark  of  various  deciduous 
and  coniferous  trees  in  oak  woods,  open  areas,  and  swamps.  Although 
the  species  was  common,  it  was  never  found  fertile. 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  Connecticut, 
New  Jersey,  North  Carolina,  Smoky  Mountains,  Michigan,  Ontario,  Min¬ 
nesota,  Black  Hills,  Arizona,  Idaho,  Washington,  British  Columbia, 
Alaska,  Saskatchewan,  Manitoba,  Baffin  Island:  Arctic-boreal  element; 
circumboreal. 

50.  PSEUDEVERNIA  Zopf 

204.  Pseudevernia  furfuracea  (L.)  Zopf,  Beih.  Bot.  Centralbl.  14: 
124.  1903.  Lichen  furfuraceus  L.  Sp.  PI.  1146.  1753. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2135  (102). 

Pseudevernia  furfuracea  is  generally  thought  of  as  a  northern  or 
high  altitude  species  (one  of  the  few  temperate  species  of  this  genus) 
found  very  commonly  in  spruce-fir  forests  on  conifers.  It  was,  therefore, 
significant  that  this  species  was  found  growing  on  a  dead  Chamaecyparis 
thyoides  in  the  cedar  bog  having  the  most  “northern”  and  oceanic  flora. 
In  this  same  bog,  I  collected  such  other  rare  (on  Long  Island)  oceanic 
and/or  northern  species  as  Loharia  pulmonaria,  L.  quercizans,  Lepto- 
gium  cyanescens,  and  Pertusaria  amara.  Pseudevernia  furfuracea  was 
also  collected  once  near  Woods  Hole  on  Cape  Cod  (Brodo  3926 )  and 
once  on  Nantucket  Island  ( Brodo  4071),  both  on  Pinus  rigida  in  oceanic 
pine-oak  forests. 

Hale  (1955c)  discussed  some  of  the  morphological  variation  of 
this  species,  as  well  as  commenting  on  its  chemistry,  especially  as  the 
species  occurs  in  North  America.  Hale  (1956b)  later  discussed,  in 
greater  detail,  its  chemical  variations  throughout  the  world,  particularly 
in  Europe. 

Distribution  —  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes-Rocky  Mountain  unit  (map:  Hale,  1955c);  Europe, 
North  Africa  (ibid). 


238  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

51.  CETRARIA  Adi. 

205.  Cetraria  ciliaris  Ach.  Lich.  Univ.  508.  1810. 

Material  seen  —  KINGS  COUNTY :  East  New  York,  G.  B.  Brain- 
erd,  (1866?)  (BKL  031917).  SUFFOLK  COUNTY:  Imshaug  25796 
(86),  25818  (86),  25829  (86);  Brodo  1067  (130),  1098  (78),  2094 
(78),  2137  (102),  2271  (87),  2493  (23),  3132  (68),  3837  (66); 
16  specimens  collected  by  Latham  (Latham). 

Cetraria  ciliaris  has  been  the  subject  of  a  detailed  population  study 
(Hale,  1963).  Hale  discussed  the  local  and  geographic  distributions  of 
the  three  chemical  races  known  within  the  species:  the  KC  +  red,  C  — 
a'ectoronic  acid  strain,  the  C  +  red  olivetoric  acid  strain,  and  KC  — , 
C  —  protolichesteric  acid  strain. 

The  two  most  common  strains  (olivetoric  and  alectoronic)  are 
both  represented  on  Long  Island.  Only  2  of  the  10  specimens  I  collected 
contained  olivetoric  acid,  and  none  of  Latham’s  16  specimens  were  of 
that  strain.  Hale  (1963)  demonstrated  that  the  distribution  of  the  two 
strains  shows  no  correlation  with  substrate,  microclimate,  or  other  envi¬ 
ronmental  factors.  Their  distribution  in  the  Appalachians  and  in  North 
America  in  general  shows  extensive  overlapping  and  there  is  no  basis 
for  giving  them  taxonomic  recognition. 

The  ecology  of  C.  ciliaris  on  Long  Island  is  almost  precisely  as  noted 
by  Hale  (1963).  It  is  usually  found  on  Finns  rigida  and  Chamaecy paris 
thyoides  in  typical  photophilous  conifer  lichen  communities,  but  on 
Long  Island  it  is  almost  entirely  restricted  to  bog  and  swamp  situations 
(figure  33).  It  is  occasionally  found  on  Primus  maritima  in  the  dune 
com  munity. 

Distribution  —  Appalachian-Great  Lakes-Rocky  Mountain  distribu¬ 
tion  with  west  coast  population  of  the  protolichesteric  acid  strain  (map: 
Hale,  1963):  Temperate  element,  North  Temperate  subelement  (?) 
(Ahti,  1964);  Europe  (Ahlner,  1940);  Asia  (Vainio,  1928). 

206.  Cetraria  fendleri  (Nyl.)  Tuck.  Gen.  Lich.  280.  1872.  Platsyma 
fendleri  Nyl-  Syn.  Lich.  1:  309.  1860. 

Material  seen  —  SUFFOLK  COUNTY:  Manorville,  Latham  7767a, 
May  28,  1937  (Latham);  Napeague,  Latham  25986 ,  March  11,  1947 
(NYS). 

This  species  is  apparently  very  rare  on  Long  Island,  although  I  col¬ 
lected  it  in  southern  New  Jersey  (Brodo  3698),  Nantucket  ( Brodo 
4112),  and  Cape  Cod  (Brodo  4191,  4312).  Latham’s  comment  on  his 
no.  25986,  “fairly  common  on  pine  bark,  trunk  and  twigs  in  barren 
sandy  grounds  .  .  .  ,”  must  have  pertained  to  a  very  local  population  in 
that  area. 

Cetraria  fendleri  is  a  typical  member  of  the  photophilous  community 
on  pine  twigs,  along  with  Parmeliopsis  placorodia.  Interestingly,  it  has  an 
almost  identical  North  American  distribution.  Culberson  (1961b)  also 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  239 

has  commented  on  the  eco'ogical  and  phytogeographic  similarities  of 
these  two  species. 

In  North  Carolina,  Cetraria  fendleri  is  an  abundant  pine-bark  lichen 
occurring  most  abundantly  in  the  Piedmont  region  (Culberson,  1958). 

Distribution  —  Temperate  element,  Appalachian  subelement,  Appa 
lachian-Great  Lakes-Rocky  Mountain  unit  (Culberson,  1961b);  endemic. 

207.  Cetraria  islandica  (L.)  Ach.  Meth.  Lich.  293.  1803.  Lichen 
islandicus  L.  Sp.  PI.  1  145.  1753. 

subsp.  crispa  (Ach.)  Cromb.  Grevillea  12:  73.  1884.  Cetraria  islandi¬ 
ca  y  C.  crispa  Ach.  Lich.  Univ.  513.  1810. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3351  (8);  Plain  Edge, 
S.  Cain ,  August  3,  1936  (NY).  SUFFOLK  COUNTY:  16  specimens 
collected  by  Imshaug  and/or  Brodo;  63  specimens  collected  by  Latham 
(Latham);  Southampton,  Chite,  September  3  to  7,  1898  (NY);  Montauk, 
Copeland  2090,  June  7,  1941  (MSC). 

Cetraria  islandica  is  a  widely  distributed  circumboreal  species  which 
includes  a  number  of  morphological  and  chemical  variants.  Imshaug 
(1957)  presented  a  detailed  discussion  of  these  variants  and  their  taxo¬ 
nomic  status.  Following  Imshaug’s  argument,  the  Long  Island  material 
—  having  only  marginal  pseudocyphellae  and  showing  a  PD  —  reaction 
in  the  medulla  —  can  be  referred  to  subspecies  crispa.  On  the  species 
level,  this  taxon  would  be  C.  ericetorum  Opiz  (Ahti,  1964). 

Roy  Latham  became  particularly  interested  in  this  species  and  pub¬ 
lished  a  series  of  five  papers  on  its  Long  Island  distribution  (Latham, 
1945,  1946,  1947,  1948).  In  these  papers,  he  noted  the  various  stations 
where  the  lichen  grew,  the  condition  and  extent  of  each  colony,  and  its 
history  as  to  hurricane  or  fire  damage.  After  much  field  work,  he  con¬ 
cluded  that,  although  C.  islandica  is  commonly  found  on  exposed  hill¬ 
tops  and  beaches,  the  species  is  found  just  as  often  “.  .  .  in  fiat  woodlands, 
locally  remote  from  hilltops  and  exposed  beaches  .  .  .”  (Latham,  1947). 

My  own  field  experience  bears  out  Latham's  observations.  Cetraria 
islandica  is  found  as  a  conspicuous  member  of  the  communities  on  sand 
dunes  and  grassy  "downs”  (p.  60),  along  with  Cladonia  submitis  and 
C.  boryi.  It  is  interesting  to  note  that  it  still  can  be  found  in  surprising 
abundance  in  central  Nassau  County,  along  the  Meadowbrook  Parkway 
on  the  remains  of  the  Hempstead  Plains,  just  as  it  was  in  1936  in  nearby 
Plain  Edge  (cf.  above)  prior  to  the  suburbanization  of  the  area. 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  Connecticut, 
Michigan,  Ontario,  Minnesota,  Rocky  Mountains  (Imshaug,  1957a), 
Washington,  Alaska,  Saskatchewan,  Manitoba,  Canadian  archipelago, 
Quebec,  Baffin  Island:  Arctic-boreal  element;  circumboreal. 

208.  Cetraria  tuckermanii  Oakes  in  Tuck.  Amer.  J.  Sci.  Arts  45:  48. 
1843. 

Material  seen  —  QUEENS  COUNTY:  Jamaica,  G.  B.  Brainerd, 
(1866?)  ( BKL  031916).  SUFFOLK  COUNTY:  North  Sea,  Latham 
35933,  May  20,  1954  (Latham). 


240  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  nomenclature  of  this  species  had  long  been  a  source  of  con¬ 
fusion  until  Imshaug  (1954)  clarified  the  identities  of  various  members 
of  the  group.  Cetraria  tuckermanii  Herre  as  treated  by  Fink  (1935) 
should  be  called  C.  herrei  Imsh.  (a  species  of  the  west  coast);  Fink’s 
C.  lacunosa  Ach.,  at  least  in  part,  is  actually  C.  tuckermanii  Oakes  in 
Tuck. 

On  Long  Island,  the  species  is  very  rare.  I  found  it  only  once  in 
southern  New  Jersey  ( Brodo  3597).  Where  it  occurs,  it  is  apparently  a 
member  of  the  bog  community  on  Chamaecy paris  with  other  bog  Cetra- 
riae  (  e.g.,  C.  ciliaris,  C.  viridis) . 

Distribution  - —  Temperate  element,  Appalachian  subelement,  Appa¬ 
lachian-Great  Lakes  unit  (Hale,  1961a);  endemic. 

209.  Cetraria  viridis  Schwein.  in  Halsey,  Ann.  Lyc.  Nat.  Hist.  N.  Y. 
1 :  16.  1824. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25789  (86), 
25801  (86),  25828  (86),  25849  (86);  Brodo  1094  (78),  2127  (102), 
2241  (87);  Flanders,  Latham ,  May  31,  1925  (Latham);  Calverton, 
Latham,  May  1,  1960  (Latham);  Riverhead,  Latham,  May  25,  1960 
(Latham);  Riverhead,  Latham,  June  17,  1960  (Latham);  Riverhead, 
Latham  2369,  June  22,  1924  (Latham);  Montauk  Point,  Latham  36972, 
September  15,  1949  (Latham). 

Cetraria  viridis  has  usually  been  considered  as  a  synonym  of  C. 
juniperina  (L.)  Ach.  (see  Fink,  1935).  However,  the  dark  yellow-green 
to  almost  grey-green  upper  surface  of  C.  viridis,  together  with  its  small, 
finely  divided  almost  lacy  margins  and  its  restricted  east  coast  distri¬ 
bution.  all  serve  to  distinguish  it  from  the  pure  yellow  (sometimes  dark 
yellow)  more  broadly  lobed,  more  northern  C.  juniperina  sens.  str. 

In  the  northeastern  coastal  plain,  C.  viridis  is  narrowly  restricted  to 
Chamaecyparis  bogs  on  the  white  cedar  trees  themselves  (figure  34),  or, 
more  rarely,  on  Pinus  rigida  or  Vaccinium  corymbosum  (p.  50).  1  have 
collected  it  in  bogs  on  Chamaecyparis  in  New  Jersey  (Brodo  3672,  3791) 
and  Cape  Cod  ( Brodo  4348) . 

Distribution  —  Massachusetts,  New  Jersey:  Temperate  element, 
Coastal  Plain  subelement;  endemic. 

52.  ANZIA  Stizenb. 

210.  Anzia  colpodes  (Ach.)  Stizenb.  Flora  45:  243.  1862.  Lichen 
colpodes  Ach.  Lich.  Suec.  Prodr.  124.  1798. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1772  (127),  1830 
(125),  1898  (114),  2496  (67),  3282  (119);  Orient  Point,  Latham, 
April  18,  1910  (NYS);  Orient,  Latham,  May  3,  1914  (Latham);  Na- 
peague,  Latham  8120  (Latham);  Napeague,  Latham  8122B,  November  6, 
1938  (Latham). 

With  its  thick,  highly  branched,  black  hypothallus,  Anzia  colpodes 
can  hardly  be  confused  with  any  other  species  on  Long  Island.  Super¬ 
ficially,  however,  it  sometimes  gives  the  appearance  of  being  a  form  of 
Hypogymnia  physodes.  It  is  treated  in  the  genus  Parmelia  by  Fink  (1935). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


241 


Anzia  colpodes  was  collected  almost  exclusively  in  humid,  oceanic 
oak  and  oak-pine  forests  of  the  eastern  tip  of  Long  Island  (figure  64), 
as  well  as  on  Nantucket  Island  ( Brodo  4128)  and  Cape  Cod  (Brodo 
4285,  4290).  It  was  always  found  on  Quercus  velutina,  usually  at  breast 
height,  although  a  few  of  Latham’s  specimens  were  from  Juniperus 
virginiana. 

Distribution  —  Eastern  United  States,  especially  in  southern  Appa¬ 
lachian  and  Ozark  Mountains  (map:  Hale,  1955c):  Temperate  element, 
Appalachian  subelement,  Appalachian-Ozark  unit  (figure  23);  Tasmania 
(Wilson,  1893  in  Wetmore,  1963). 

USNEACEAE 

53.  EVERNIA  Ach. 

211.  Evernia  mesomorpha  Nyl.  Lich.  Scand.  74.  1861. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  692  B  (81),  2095 
(78). 

This  species,  included  under  Evernia  prunastri  (L. )  Ach.  var.  tham- 
nodes  Flot.  by  Fink  (1935),  is  characterized  by  relatively  soft,  flexible, 
highly  irregular  and  angular,  sorediate  lacinae.  It  is  very  rare  on  Long 
Island,  occurring  on  trees  and  shrubs  in  cedar  bogs.  It  was  also  found 
on  pines  ( Finns  rigida)  in  a  pine-oak  forest  on  Nantucket  Island  ( Brodo 
4076),  and  on  open  downs  and  in  forests  on  Cape  Cod  (Brodo  4175, 
4314,  4495). 

Distribution  —  Nova  Scotia,  Maine,  Massachusetts,  Connecticut, 
Michigan.  Black  Hills,  Saskatchewan,  Manitoba,  Ontario:  Temperate 
element,  North  Temperate  subelement  (?);  listed  as  having  an  Appa¬ 
lachian-Great  Lakes  distribution  by  Hale  (1961a);  Europe  (Poelt, 
1963);  Asia  (Zahlbruckner,  1930). 

54.  ALECTORIA  Ach. 

212.  Alectoria  glabra  Mot.  Fragm.  FI.  Geobot.  6(3):  448.  1960. 

Material  seen  —  SUFFOLK  COUNTY:  Patchogue,  Latham,  June 

11,  1921  (Latham). 

The  Long  Island  specimen  was  compared  with  an  isotype  of  A. 
glabra  (herb.  US)  and  the  two  specimens  agreed  in  all  characteristics 
except,  perhaps,  the  general  color,  which  was  somewhat  paler  in  the  type. 
Both  showed  the  abundant  non-isidiate  soralia  and  the  PD  +  red  re¬ 
action.  Alectoria  americana  Motyka,  which  is  the  more  common  North 
American  member  of  the  A.  jubata- complex,  is  PD  —  and  lacks  soredia 
entirely.  These  species  are  discussed  more  thoroughly  by  Motyka  (1964). 

Alectoria  glabra  was  most  likely  considered  under  the  name  A.  jubata 
(L.)  Ach.  in  Fink’s  (1935)  flora. 

There  is  some  question  as  to  whether  the  Latham  specimen  of 
A.  glabra  actually  was  collected  on  Long  Island.  It  is  possible  that  it 
was  collected  elsewhere  in  North  America,  was  sent  to  Latham  on  ex¬ 
change,  and  somehow  became  mislabeled  (as  was  the  case  with  a  few 


242  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

specimens  from  the  Pacific  northwest  area) .  Until  this  basically  northern 
species  is  collected  again  on  Long  Island,  or  in  the  Cape  Cod  region 
which  has  a  more  northern  flora,  its  presence  on  Long  Island  must 
remain  questionable. 

Distribution  —  Washington  (type  locality),  Rocky  Mountains  from 
British  Columbia  to  Colorado,  Ontario  and  Newfoundland  (Motyka, 
1964):  Temperate  element,  North  Temperate  subelement;  endemic. 

213.  Alectoria  nidulifera  Norrl.  in  Nyl.  Flora  58:  8.  1875. 

Material  seen  —  QUEENS  COUNTY:  Jamaica,  G.  B.  Brainerd, 

1866  (BKL).  SUFFOLK  COUNTY:  17  specimens  collected  by  Imshaug 
and/or  Brodo;  11  specimens  collected  by  Latham  (Latham). 

This  species,  which  Fink  (1935)  probably  considered  under  the 
name  A.  chalybeiformis  (L.)  Rohl.,  is  a  frequent  member  of  the  pine 
bark  community.  It  is  found  in  open  pine  barrens,  pine  forests,  and  bogs, 
mostly  on  Pinus  rigida  and  Chamaecy  parts  thyoides  (figure  40).  Occa¬ 
sionally  it  is  collected  from  dead  twigs  or  tangled  stumps  of  Hudsonia 
tomentosa  close  to  the  ground  in  open  sand  barrens. 

Motyka  ( 1964)  gives  a  detailed  description  of  the  species  and  points 
out  a  number  of  differences  between  the  American  and  European  popu¬ 
lations. 

Distribution  —  Quebec,  northeastern  United  States  south  to  Vir¬ 
ginia  (Motyka,  1964);  Nova  Scotia,  Maine,  Massachusetts,  Connecticut, 
North  Carolina,  Tennessee,  Michigan,  Wisconsin,  Arizona,  British  Colum¬ 
bia,  Saskatchewan,  Ontario:  Temperate  element,  Appalachian  subelement, 
Appalachian-Great  Lakes  unit  (Hale,  1961a);  Europe  (Motyka,  1964); 
Asia  (Vainio,  1928). 

55.  RAMALINA  Ach. 

214.  Ramalina  complanata  (Sw.  in  Ach.)  Ach.  Lich.  Univ.  599. 
1810.  Lichen  complanatus  Sw.  in  Ach.  Kgl.  Vet.  Akad.  Nya  Handl. 
290.  1797. 

Material  seen  —  Orient,  Latham,  April  20,  1920  (Latham). 

The  specimen  upon  which  this  record  is  based  is  sterile  and  poorly 
developed.  In  view  of  the  species’  normally  southern  or  tropical  distri¬ 
bution,  such  a  record  will  have  to  be  viewed  with  some  skepticism,  at 
least  until  more  material  is  collected  in  the  area. 

Material  of  R.  complanata  from  the  Howe  collection  at  the  Farlow 
Herbarium  was  compared  with  Latham’s  collection.  Except  for  its  being 
sterile,  Latham’s  material  agreed  well  with  a  specimen  from  Corpus 
Christi,  Texas  ( Howe  2553),  as  well  as  one  from  Lake  Ngunga,  British 
East  Africa  (Howe  1786,  S.  M.  Allen,  August  27,  1909).  All  had  broad, 
heavy,  stiff,  more  or  less  striate  and  rimose  lacinae  with  conspicuous 
white  pseudocyphellae  or  tuberculae,  and  all  had  PD  — .  KOH  —  medul¬ 
lary  reactions. 

The  species  was  described  from  Jamaica,  and  according  to  Howe 
(  1914)  it  is  “common  in  the  Austral  Zone.”  If  the  Long  Island  specimen 
is  correctly  identified,  it  would  not  he  the  first  example  of  a  tropical 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  243 

species  which  has  migrated  up  the  Atlantic  coastal  plain  as  far  north 
as  Long  Island  (see  Cladonia  evansii).  The  Latham  specimen  was  found 
on  J  uni  perns. 

Distribution  —  Florida,  Texas,  West  Indies:  Tropical  element. 
Coastal  Plain  subelement  (map:  Howe,  1914);  East  Africa  (above). 

215.  Ramalina  fastigiata  (Lilj.)  Ach.  Lich.  Univ.  603.  1810.  Lichen 
calcaris  var.  fastigiata  Lilj.  Utkast  Svensk.  FI.  426.  1792. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25766  (121); 
Brodo  692A  (81),  1024  (112),  1727A  (131),  1732  (131),  1816  (125), 
1956  (85),  2962A  (95),  3307  (129);  Orient,  Latham ,  April  20,  1920 
(Latham);  Orient,  Latham  42,  May  23,  1914  (Latham);  Sag  Harbor, 
Britton  213,  July  17,  1898  (NY);  Sag  Harbor,  Britton,  July  13,  1897 
(NY). 

Ramalina  fastigiata  is  a  variable  species  characterized  by  its  small, 
straight  ellipsoid  spores,  and  broad,  usually  short,  often  somewhat 
channeled  lacinae.  Magnusson  apparently  believed  the  species  should  be 
greatly  subdivided,  and  he  had  annotated  specimens  from  many  Ameri¬ 
can  herbaria  with  unpublished  names  such  as  R.  americana  and  R. 
confusa  Magn.  I  have  studied  material  annotated  by  Magnusson  as 
americana,  confusa,  and  fastigiata,  and  can  find  no  constant  character 
or  combination  of  characters  to  warrant  the  recognition  of  more  than 
one  species. 

The  species  occurs  in  the  oceanic  eastern  tip  of  Long  Island,  mainly 
in  the  exposed  lee  dune  and  down  communities  or  in  well-lighted  forests 
growing  on  various  deciduous  trees  and  shrubs. 

Distribution  —  Throughout  eastern  United  States  (map:  Howe, 
1914):  Temperate  element,  East  Temperate  subelement;  Europe;  Asia 
(Zahlbruckner,  1930;  Vainio,  1928). 

216.  Ramalina  stenospora  Mull .  Arg.  Flora  60:  477.  1877. 

Material  seen  —  SUFFOLK  COUNTY:  Orient,  Latham,  October  1, 

1914  (Latham);  Orient,  Latham,  April  18.  1923  (Latham);  Orient 
Point,  Latham,  November  26,  1909  (NYS);  Orient,  Latham  742,  Octo¬ 
ber  5,  1918  (NYS):  Southampton,  Morgan  (Howe  1677)  September  15, 
1909  (FH:  Howe);  Southampton,  Carnegie  ( Howe  2659),  June  22, 
1913  (FH:  Howe);  Southampton,  Carnegie  (Howe,  Lich.  Nov.  Angl. 
64),  August  20,  1914  (FH:  Howe,  MSC). 

Although  R.  stenospora  has  been  collected  in  eastern  Long  Island 
a  number  of  times,  I  myself  have  never  seen  it  in  the  field.  It  is  basically 
a  southern  species  closely  related  to  R.  montagnei  De  Not.,  which  has 
distinctly  terete  or  subterete  rather  than  strap-shaped  lacinae.  Howe 
(1914)  reported  R.  montagnei  from  Jamaica,  Cuba,  Louisiana,  and 
Florida.  I  have  also  seen  a  specimen  from  the  Bahama  Islands. 

Ramalina  stenospora  appears  to  be  a  member  of  the  community 
on  coastal  Juniperus  virginiana,  along  with  R.  willeyi. 


244 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Distribution  —  West  Indies,  Gulf  and  Atlantic  coasts  north  to 
Massachusetts  (map:  Howe,  1914):  Temperate  element,  Coastal  Plain 
subelement:  endemic. 

217.  Ramalina  willeyi  R.  H.  Howe,  Bryol.  17:  36.  1914. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25764  (121); 
Brodo  2834  (115),  2962 B  (95);  Orient,  Latham ,  April  20,  1920  (La¬ 
tham);  Napeague,  Latham,  November  6,  1938  (Latham);  Orient,  Latham 
7436,  June  5,  1933  (Latham);  Orient,  Latham  8576,  June  5,  1933 
(Latham);  Orient,  Latham  8585,  April  30,  1939  (Latham);  Montauk, 
Hither  Beach,  Latham  24010,  October  28,  1945  (Latham);  Orient  Point, 
Latham,  December  20,  1909  (NYS);  Orient  Point,  Latham,  May  2,  1910 
(NYS);  Promised  Land,  Latham,  January  21,  1947  (NYS). 

Rat)ialina  willeyi,  with  its  subterete  lacinae  and  KOH  +  red  medul¬ 
lary  reaction,  is  easily  identified.  Although  the  type  could  not  be  found 
in  the  Howe  herbarium,  an  isotope  from  the  Willey  herbarium  at  the 
Smithsonian  was  examined  by  Hale,  who  reports  (pers.  comm.)  that 
the  KOH  +  constituent  is  salacinic  acid.  Howe  (1914)  seemed  to  re¬ 
gard  the  species  as  basically  KOH  — ,  helping  to  distinguish  it  from  the 
West  Indian  species  R.  attennata,  which  he  said  was  KOH  +.  However, 
he  stated  that  he  had  seen  specimens  of  R.  willeyi  with  a  distinct  colora¬ 
tion  in  KOH.  All  the  Long  Island  specimens  contain  salacinic  acid.  One 
specimen  from  Cape  Cod  ( Brodo  4378),  tentatively  identified  as  R. 
willeyi,  contains  protocetraric  acid  (by  chromatography).  A  KOH — 
Florida  specimen  (in  herb  MSC),  annotated  by  Magnusson  as  R.  willeyi, 
had  the  flattened  lacinae  and  slightly  curved  spores  of  R.  complanata.  It 
would,  therefore,  seem  that  salacinic  acid  is  almost  a  constant  component 
of  the  species,  with  protocetraric  acid  being  a  rare  alternate. 

Ramalina  willeyi  is  a  member  of  the  aerohaline  community  on 
Juniperus  virginiana.  The  species  was  found  to  be  more  luxuriant  and 
more  common  on  Nantucket  Island  and  on  Cape  Cod. 

Distribution  —  All  along  the  Gulf  and  Atlantic  coasts  north  to 
Cape  Cod  (map:  Howe,  1914):  Temperate  element.  Coastal  Plain  sub¬ 
element;  endemic. 


56.  USNEA  P.  Br.  ex  Adans. 

218.  Usnea  longissima  Ach.  Lich.  Univ.  626.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Napeague,  Latham,  May 
30,  1922  (Latham);  Northwest,  Latham,  May  18,  1949  (Latham). 

This  species,  like  Alectoria  glabra,  is  characteristic  of  the  spruce-fir 
forests  of  the  north.  I  did  not  collect  it  anywhere  on  Cape  Cod  or 
Nantucket  Island,  where  other  species  of  Usnea  were  abundant,  nor  in 
any  of  the  localities  on  Long  Island  having  “northern”  floras.  However, 
according  to  the  specimens  cited  by  Motyka  (1936-38)  in  his  mono¬ 
graph,  the  species  has  a  distribution  which  could  conceivably  include 
Long  Island.  It  is  also  quite  possible  that  the  specimens,  like  some 
others  in  the  Latham  collection,  were  mislabeled. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  245 

Distribution  —  Nova  Scotia,  Michigan,  Ontario,  Minnesota,  Wash¬ 
ington,  Alaska:  Temperate  element,  North  Temperate  subelement  (Ahti, 
1964);  Europe;  Asia  (Asahina,  1956). 

219.  Usnea  mutabilis  Stirt.  Scot.  Natural.  6:  107.  1881. 

Material  seen  —  Orient,  Latham,  May  23,  1914  (Latham);  Orient, 
Latham  8612B,  April  30,  1925  (Latham);  Orient,  Latham  8613,  April 
30,  1925  (Latham). 

The  densely  isidiate  branches  and  red  medulla  of  this  species  quickly 
separate  it  from  all  other  Long  Island  Usneae.  Link  (1935)  included 
U.  mutabilis  as  a  synonym  of  U.  florida,  an  entirely  different  species. 

Usnea  mutabilis,  having  been  collected  on  Long  Island  only  by  Roy 
Latham,  is  one  of  the  rarest  of  the  Long  Island  lichens,  and  may  in  fact 
be  “extinct”  on  the  island  at  the  present  time  (cf.  p.  276).  All  three 
Latham  collections  came  from  Orient  prior  to  the  1938  hurricane  which 
devastated  so  much  of  that  area  and  washed  away  so  many  rare  species 
of  lichens  (p.  276;  Latham,  1945).  On  nearby  Nantucket  Island  and 
Cape  Cod,  the  species  still  grows  luxuriantly  in  some  localities,  particu¬ 
larly  in  pine-oak  forests  and  bogs  on  pines  or  other  trees. 

Distribution  —  Throughout  eastern  United  States,  especially  in  the 
south  (Motyka,  1936-38):  Temperate  element.  East  Temperate  sub¬ 
element;  endemic. 

220.  Usnea  strigosa  (Ach.)  A.  Eaton,  Man.  Bot.  ed.  5,  431.  1829. 
Usnea  florida  y.  U.  strigosa  Ach.  Meth.  Lich.  310,  pi.  6,  f.  3.  1803. 

Material  seen  —  (Medulla  red)  QUEENS  COUNTY:  Ridgewood, 
G.  B.  Brainerd ,  1866  (BKL  031932).  SULLOLK  COUNTY:  39  speci¬ 
mens  collected  by  Imshaug  and/or  Brodo;  19  specimens  collected  by 
Latham  (Latham);  Riverhead,  Peck  (NYS);  Sayville,  Lloyd  137, 
December  2,  1896  (NY). 

(Medulla  white)  QUEENS  COUNTY:  Ridgewood,  G.  B.  Brainerd, 
1866  (BKL).  SULLOLK  COUNTY:  15  specimens  collected  by  Imshaug 
and/or  Brodo;  Mattituck,  Latham,  July  4,  1914  (Latham);  Montauk,  La¬ 
tham,  May  6,  1920  (Latham);  Montauk,  Latham,  April  17,  1934 
(Latham);  Shelter  Island,  Swamp  N.  of  —  (?),  Latham  22221,  October 
26,  1944  (Latham);  Riverhead,  Latham  36871 B,  3687 1C,  May  16,  1960 
(Latham);  Three  Mile  Harbor,  Latham  34091 B.  April  17,  1947  (La¬ 
tham);  Quogue,  Latham  34313,  September  2,  1950  (Latham). 

Of  all  the  difficult  groups  in  the  difficult  genus  Usnea,  the  U.  barbata 
group  is  certainly  one  of  the  most  challenging.  Hale  (1962)  recently 
pointed  out  that  U.  strigosa  is  made  up  of  a  number  of  chemical  strains, 
among  them,  a  norstictic  acid  positive  strain  and  a  norstictic-less  strain, 
with  red  medullary  color  having  no  taxonomic  value.  Henry  Imshaug 
and  I,  working  independently  from  Hale,  arrived  at  precisely  the  same 
conclusions. 

Of  the  50  specimens  of  U.  strigosa  having  a  red  medulla  that  we 
chromatogrammed,  30  (60  percent)  contained  norstictic  acid,  and  20 


246  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

(40  percent)  lacked  norstictic  acid.  All  specimens  with  a  white  medulla 
that  were  tested  (19)  contained  norstictic  acid.  The  psoromic  acid  strain 
reported  by  Hale  (1962)  was  not  represented  at  all.  Two  Latham  speci¬ 
mens  (Montauk,  April  17,  1934;  Montauk,  May  6,  1920)  contained 
salacinic  acid  and  would  he  referable  to  either  the  U.  arizonica  Motyka 
population  or  perhaps  to  the  population  represented  by  the  type  of 
U.  subfusca  Stirt.  (which  is  also  a  member  of  the  strigosa  complex) 
(see  Hale,  1962). 

Thus  circumscribed,  U.  strigosa  becomes  fairly  easy  to  identify, 
being  a  shrubby,  densely  strigose  species.  The  number  of  apothecia  (none 
to  many),  color  of  the  medulla  (pure  white  to  dark  rusty  red  with  all 
intermediates),  and  the  presence  of  norstictic  acid  all  are  variable.  It 
should  be  noted  that  the  concentration  of  norstictic  acid  varies  within 
the  thallus  as  well.  In  the  white  medulla  form  a  clear  KOH  +  red 
reaction  often  could  be  seen  only  in  the  medulla  of  the  apothecia,  with 
the  medulla  of  the  filaments  being  perfectly  negative  with  KOH,  or 
at  best,  pa'e  yellow. 

In  U.  strigosa,  we  once  again  see  the  norstictic  - —  psoromic 
( — salacinic)  shift  which  occurs  so  often  in  closely  related  lichens. 

Usnea  strigosa  is  fairly  common  in  well-lighted  oak  woods,  bogs, 
lee  dune  thickets,  etc.,  throughout  eastern  Long  Island.  It  occurs  exclu¬ 
sively  on  deciduous  trees  and  shrubs. 

Distribution  — -  Throughout  eastern  United  States  (map;  Hale, 
1962):  Temperate  element,  East  Temperate  subelement;  Asia  (Zahl- 
bruckner,  1930). 

221.  Usnea  trichodea  Ach.  Meth.  Lich.  312.  p.  8,  f.  1.  1803. 

Material  seen  - —  SUFFOLK  COUNTY:  hnshaug  25802  (86), 

25811  (86);  Brodo  897  (56),  1026  (112),  1028  (112),  1038  (112), 
1666  (88),  2159  (102),  2161  (102),  2247 A  (87);  13  specimens  col¬ 
lected  by  Latham  (Latham);  Orient,  Young  (BKL);  Orient,  Latham, 
December  20,  1909  (NYS);  Brookhaven,  Ames,  May  1910  (NYS). 

The  long,  very  slender,  articulated  filaments  of  U.  trichodea  are  a 
common  feature  of  the  corticolous  bog  lichen  communities  on  all  species 
of  trees  (figure  39). 

Distribution  —  Nova  Scotia  (type  locality),  east  coast  south  to 
Florida  and  Texas  (see  Motyka,  1936-38),  coastal  British  Columbia 
(Weber  and  Shushan,  1959):  Temperate  element,  Coastal  Plain  sub¬ 
element  (?);  Asia  (Zahlbruckner,  1930).  Asahina  (1956)  states  that 
U.  hossei  Vain.  f.  suhtrichodea  Asahina  from  Japan  has  often  been  con¬ 
fused  with  U.  trichodea,  and  Zahlhruckner’s  report  from  China  may 
represent  a  misidentification.  The  B.C.  report  is  from  the  Queen 
Charlotte  Islands,  known  for  many  Asian  disjuncts,  and  may  also  repre¬ 
sent  U.  hossei.  I  have  not  examined  the  specimen. 

222.  Usnea  sp.  (Usnea  subfusca  sensu  Mot.) 

Material  seen  —  SUFFOLK  COUNTY:  hnshaug  25800  (86), 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


247 


25822  (86);  Brodo  1820  (125)  (?),  2104  (86),  2106A  (86),  2107  (86), 
2134  (102),  2247B  (87),  2262  (87),  2269  (87),  2803  (102);  North¬ 
west,  Latham  27209,  April  18,  1947  (Latham);  Montauk,  SW  of  —  (?), 
Fort  Pond,  Latham  34082,  July  8,  1957  (Latham);  Riverhead,  North 
Swamp,  Latham  36877,  May  25,  1960  (Latham);  Riverhead,  Latham 
36946,  June  17,  1960  (Latham). 

Most  Long  Island  specimens  of  this  species  were  identified  by  Herre 
as  U.  subfusca  Stirt.  (often  as  var.  halei  Herre)  and,  indeed,  they  agree 
in  most  respects  with  the  description  of  U.  subfusca  given  hy  Motyka 
(1936-38)  in  his  monograph.  However,  as  Hale  (1962)  pointed  out, 
Stirton’s  specimen  of  U.  subfusca  (which  1  have  also  examined  both 
morphologically  and  chemically)  contains  salacinic  acid  and  has  the 
strigose  habit  of  a  member  of  the  Usnea  strigosa  complex.  This  leaves 
the  species  referred  to  by  Motyka  as  U.  subfusca  without  a  name.  Since 
there  are  several  species  cited  (or  even  described  as  new)  by  Motyka 
which  seem  to  be  close  to  his  U.  subfusca,  it  is  probable  that  one  of 
these  names  could  be  applied  here,  and  so  no  nomen  novum  is  provided 
at  this  time.  A  likely  candidate  is  U.  merrillii  Motyka,  and  Herre  even 
annotated  some  specimens  of  this  species  under  that  name.  However,  the 
exsiccats  cited  by  Motyka  as  U.  merrillii  (Merrill,  Lich.  64,  99,  and  130), 
which  I  examined  at  the  University  of  Michigan  herbarium  were  dif¬ 
ferent  from  one  another,  and,  in  one  case  (no.  99),  was  a  mixed  collec¬ 
tion.  Confirmation  must  await  an  examination  of  the  holotype  of  this 
and  other  suspect  species. 

The  Long  Island  material  is  very  variable  in  many  respects.  Although 
its  general  aspect  is  dark  ashy  green  to  olivaceous,  it  sometimes  becomes 
stramineous  toward  the  younger  branchlets  and  is  often  mottled  yellowish 
and  grey-green  in  younger  portions.  Branching  is  loose,  with  irregular 
side  branches  and  common  dichotomies,  although  the  end  shoots  fre¬ 
quently  have  few  or  no  side  branches.  Papillae  and  sometimes  small 
tubercles  usually  cover  the  main  branches,  but  sometimes  they  are  sparse. 
Isidiate  soralia  are  generally  conspicuous  (rarely  absent)  on  the  younger 
portions  of  the  thallus.  Apothecia,  which  are  very  infrequent,  are  heavily 
pruinose,  small  (1.5-3  mm)  and  have  long  marginal  “cilia”  1-3  times 
the  diameter  of  the  disk.  The  cortex,  however,  is  always  thick  and  chon- 
droid,  and  the  medulla  is  always  white  and  thin. 

Chromatographic  analysis  showed  that  protocetraric  acid  is  present 
in  all  specimens  except  for  one  Long  Island  collection  ( Brodo  2247B), 
which  contains  barbatic  acid,  and  two  Cape  Cod  collections  ( Brodo 
4161,  4338)  which  showed  the  presence  of  fumarprotocetraric  acid. 

The  species  is  found  in  well-lighted  areas  of  bogs  and  swamps, 
being  rare  elsewhere.  It  occurs  on  all  types  of  trees,  especially  Chamae- 
cyparis  thyoides,  and  thus  is  usually  associated  with  Usnea  trichodea. 

Distribution  —  Probably  widespread  in  east  temperate  North 
America. 


248  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

TELOSCHISTACEAE 

57.  CALOPLACA  Th.  Fr, 

223.  Caloplaca  auranfiaea  (Lightf.)  Th.  Fr.  Nova  Acta  Reg.  Soc. 
Sci.  Upsal.  III.  3:  219.  1861.  (Lich.  Arct.  119.  1860).  Lichen  auran- 
tiacus  Lightf.  FI.  Scot.  2:  810.  1777. 

Material  seen  —  SUFFOLK  COUNTY.  Brodo  3311  (129),  3439 
(134). 

Both  Long  Island  specimens  assigned  to  this  species  proved  to  be 
difficult  to  identify.  In  the  case  of  Brodo  3311,  found  on  a  windswept 
Carya  on  the  Montauk  downs,  the  thallus  was  very  thin  and  almost 
devoid  of  the  yellow  tint  characteristic  of  the  species.  In  the  case  of 
Brodo  3439 ,  the  substrate  (concrete)  did  not  agree  with  Rudolph’s 
(1955)  view  that  the  species  is  found  only  on  wood  and  bark.  The 
spore,  thallus,  and  apothecial  characters  of  the  latter  specimen  seem 
to  fit  the  descriptions  of  C.  aurantiaca  very  well,  and  other  modern 
authors  (Erichsen,  1957;  Hillmann  &  Grummann,  1957;  Bertsch,  1964) 
state  that  the  species  can  be  saxicolous. 

Distribution  —  Nova  Scotia,  Connecticut,  Oklahoma,  Wisconsin, 
Arizona,  Idaho,  Washington:  Temperate  element,  North  Temperate  sub- 
element;  Europe;  Asia  (Zahlbruckner,  1930,  and  others). 

224.  Caloplaca  camptidia  (Tuck.)  Zahlbr.  Cat.  Lich.  Univ.  7:  83. 
1930.  Lecanora  camptidia  Tuck.  Proc.  Amer.  Acad.  Arts  Sci.  5:  403. 
1862. 

Material  seen  —  SUFFOLK  COUNTY:  lmshaug  25595  (52); 
Brodo  1765B  (127),  1769A  (127),  1897  (114),  2683B  (110),  3085B 
(128),  3095  (122),  3103 B  (122);  Orient,  Latham ,  May  3,  1914  (La¬ 
tham);  Greenport,  Latham  22259,  March  29,  1914  (Latham);  Orient 
Point,  Latham  6,  November  21,  1910  (NYS). 

This  is  the  only  Caloplaca  on  Long  Island  entirely  without  anthra- 
quinone  pigments.  Superficially,  it  looks  much  like  a  Lecanora  with 
pruinose,  reddish  brown  apothecia.  Its  hyaline  polarilocular  spores,  how¬ 
ever,  clearly  identify  it  as  a  Caloplaca. 

Caloplaca  camptidia  is  found  only  on  oak  bark,  and  it  is  an  occa¬ 
sional  member  of  the  breast  height  communities  on  oak  (figure  66). 

Distribution  —  Oklahoma,  Appalachians  (Rudolph,  1955):  Tem¬ 
perate  element,  Appalachian  subelement,  Appalachian-Ozark  unit  (?); 
West  Indies  (Tuckerman,  1872);  endemic. 

225.  Caloplaca  cerina  (Ehrh.  in  Hoffm.)  Th.  Fr.  Nova  Acta  Reg. 
Soc.  Sci.  Upsal.  III.  3:  218.  1861  (Lich.  Arct.  118.  1860).  Lichen 
cerinus  Ehrh.  in  Hoffm.  PI.  Lich.  2:  62,  pi.  21,  f.  13,  1789. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3311  (129);  Three 
Mile  Harbor,  East  Hampton,  Latham  2644,  April  20,  1926  (MICH). 

As  stated  by  Rudolph  ( 1955),  the  relationship  between  C.  cerina  and 
the  very  similar  C.  pyracea  is  not  always  clear  and  it  appears,  at  least 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  249 

on  Long  Island,  that  the  two  are  very  difficult  to  separate.  Typically, 
C.  cerina  has  a  prominent,  persistent,  blue-grey,  pseudothalline  margin 
(referred  to  as  the  “amphithecium”  by  Rudolph,  1955).  In  C.  pyracea , 
the  pseudothalline  margin  almost  always  is  entirely  lacking  in  the  mature 
apothecia.  Sometimes,  however,  this  grey  margin  will  begin  to  disappear 
in  C.  cerina  or  will  be  relatively  persistent  in  C.  pyracea.  (Imshaug,  1957c, 
gives  a  discussion  on  the  relationship  between  the  amphithecium  and 
the  pseudothalline  margin.)  In  these  cases,  one  must  rely  more  heavily 
on  disk  color  (which  actually  reflects  a  difference  in  the  anthraquinone 
complements  of  the  two  species;  Burgess,  in  press).  Caloplaca  cerina  has 
dusky  yellow  to  yellow-orange  disks,  whereas  C.  pyracea  has  orange 
to  red-orange  disks.  Rudolph  (1955)  states  that  there  is  a  difference  in 
hypothecial  height,  but  1  have  not  been  able  to  verify  this  in  the  material 
I  have  studied. 

Although  C.  cerina  is  a  very  rare  species  on  Long  Island,  it  is 
widely  distributed  on  a  variety  of  “calcareous”  or  “nitrogenous”  sub¬ 
strates  (cf.  p.  30).  One  specimen  (Latham  2644)  was  collected  on  a 
turtle  shell  and  the  other  found  on  an  exposed  Carya  on  the  Montauk 
downs. 

Distribution  —  Connecticut,  Michigan,  Indiana,  Wisconsin,  Arizona, 
New  Mexico,  Black  Hills,  Washington,  Saskatchewan,  Manitoba:  Tem¬ 
perate  element  (?),  North  Temperate  subelement;  Europe;  Asia  (Zahl- 
bruckner,  1930). 

226.  Caloplaca  citrina  (Hoffm.)  Th.  Fr.  Nova  Acta  Reg.  Soc.  Sci. 
Upsal.  III.  3:  218.  1861  (Lich.  Arct.  118.  1860).  Verrucaria  citrina 
Hoffm.  Deutschl.  FI.  2:  198.  1796. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25605  (E  of 
106);  Brodo  2501  (61),  2800  (84),  2828A  (115),  2828B  (115),  2841 
(115);  near  Orient  (Narrows),  Latham,  (no  date)  (Latham). 

Caloplaca  citrina  is  recognized  by  its  yellow-orange,  granular  soredi- 
ate  thallus.  The  thallus,  however,  can  vary  from  almost  an  entirely 
effuse-subareolate  condition  to  one  which  is  sterile,  thick,  and  areolate 
with  mere  traces  of  granular  soredia.  The  more  abundantly  sorediate 
specimens  had  many  small  apothecia  with  margins  commonly  becoming 
sorediate. 

The  species  apparently  is  narrowly  restricted  to  concrete  and 
mortar  substrates  and  is  a  member  of  the  aerohaline  community  (p.  59). 
On  Long  Island,  it  is  confined  to  the  coastline  (figure  77)  although 
the  species  as  a  whole  is  continental. 

Distribution  - —  South  Carolina.  Kansas,  Iowa,  Missouri,  Minnesota 
(Rudolph,  1955);  Connecticut,  Michigan,  Black  Hills,  Washington:  Tem¬ 
perate  element.  North  Temperate  subelement;  Europe;  Asia  (Zahl- 
bruckner,  1930). 

227.  Caloplaca  discolor  (Will,  in  Tuck.)  Fink,  Lich.  FI.  U.  S.  357. 
1935.  Placodium  ferrugineum  var.  discolor  Will,  in  Tuck.  Syn.  N.  Amer. 
Lich.  1  :  178.  1882. 


250  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1762  (127);  Orient 
Point,  Latham,  April  18,  1910  (NYS) ;  Orient,  Latham  22261,  March 
20,  1921  (Latham);  Orient,  Latham  22262,  May  23,  1914  (Latham); 
Orient,  Latham  1082  (22265),  May  20,  1916  (Latham). 

Caloplaca  discolor  was  placed  in  the  genus  Blastema  by  Rudolph 
(1955)  because  of  its  frequently  alga-less  margin.  Intermediates  are 
common,  however,  and  would  seem  to  indicate  that  the  species  should 
be  retained  in  Caloplaca. 

The  species  was  found  on  the  bark  of  oak  and  red  cedar  near 
the  eastern  tip  of  Long  Island. 

Distribution  —  Massachusetts,  Michigan,  Black  Hills;  endemic. 

228.  Caloplaca  feraeissima  Magn.  Bot.  Not.  2:  189.  1953. 

Material  seen  -  SUFFOLK  COUNTY:  Brodo  59-36  (53),  1825 

(125),  3919  (54). 

This  species,  described  from  a  specimen  on  concrete  from  Wis¬ 
consin,  was  found  to  be  comparatively  common  on  inland  concrete 
substrates  on  Long  Island,  although  only  three  specimens  were  collected. 
Concrete  and  mortar  closer  to  the  coast  commonly  had  C.  citrina  in  its 
place.  The  dark  orange  to  orange-brown  apothecia  usually  subtended 
by  a  trace  of  black  prothallus,  the  yellowish,  almost  totally  absent 
thallus,  and  the  narrow  spore  isthmi  combine  to  make  this  species 
rather  distinctive  and  easily  identified. 

I  have  seen  specimens  from  Michigan  and  central  New  York 
State,  and  it  is  apparently  much  more  common  than  indicated  by  the  few 
reports.  Since  it  grows  well  on  concrete  sidewalks  and  foundations,  even 
close  to  industrial  centers,  it  will  almost  surely  become  more  abundant 
in  the  future. 

Distribution  —  Central  New  York,  Michigan,  Wisconsin  (type 
locality);  endemic. 

229.  Caloplaca  f lavovirescens  (Wulf. )  Dalla  Torre  &  Sarnth.  Flecht. 
Tirol  180.  1902.  Lichen  flavovirescens  Wulf.  Schrift.  Ges.  nat.  Freunde 
Berk  8:  122.  1787. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2824  (66),  3918 
(54),  3920  (54);  Orient,  Latham  2873  (22264),  February  7,  1926 

(Latham). 

This  species  was  found  on  partially  shaded  or  exposed  concrete. 
Roy  Latham’s  specimen  from  bone  (above)  was  not  entirely  typical  of 
the  species  in  that  it  had  a  very  inconspicuous  thallus. 

Distribution  —  Nova  Scotia,  Oklahoma,  Arizona,  Black  Hills,  Sas¬ 
katchewan;  “cosmopolitan”  according  to  Rudolph  (1955):  Temperate 
element.  North  Temperate  subelement;  Europe;  Asia  ( Zahlbruckner, 
1930). 

230.  Caloplaca  pyracea  (Ach.)  Th.  Fr.  Kgl.  Svensk.  Vet.  Akad. 
Hand!.  7(2):  25.  1867.  Parmelia  cerina  var.  pyracea  Ach.  Meth.  Lich. 
176.  1803. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


251 


Material  seen  —  SUFFOLK  COUNTY:  Brodo  2963  (95);  Orient, 
Latham,  May  27,  1914  (Latham);  Greenport,  Latham  50,  May  10, 
1914  (Latham);  Orient  Point,  Latham,  April  4,  1910  (NYS);  Orient, 
Latham  3939,  March  27,  1927  (MICH);  Three  Mile  Harbor,  East  Hamp¬ 
ton,  Latham  2644  (p.p.),  April  20,  1926  (MICH). 

Caloplaca  pyracea  is  the  common  corticolous  Caloplaca  on  Long 
Island.  As  mentioned  before  (p.  248),  it  is  easily  confused  with  C.  cerina. 

Distribution  —  Nova  Scotia,  Maine,  Connecticut,  Michigan,  Indi¬ 
ana,  Arizona,  Black  Hills,  Washington,  Baffin  Island:  Arctic-boreal 
element;  northeast  Greenland  (Lynge,  1940);  Europe;  Asia  (Zahlbruck- 
ner,  1930). 


58.  XANTHORIA  (Fr.)  Th.  Fr. 

231.  Xanthoria  fallax  (Hepp  in  Arn.)  Arn.  Verh.  Zool.  Bot.  Ges. 
Wien  30:  121.  1880.  Physcia  fallax  Hepp  in  Arn.  Flora  41:  307.  1858. 

Material  seen  —  QUEENS  COUNTY:  W.  Flushing,  ( Brainerd ?) 
(BKL);  W.  Flushing  G.  B.  Brainerd  ( BKL  032033);  West  Flushing, 
(Brainerd!) ,  April  12,  1868  (BKL  032034).  SUFFOLK  COUNTY: 
Imshaug  25640  (64);  Brodo  884  (55),  1071  (98),  2118  (84),  2592 
(97),  2701  (SE  of  107),  2825  (115),  3097  (122),  3197  (32),  3359 
(S  of  97),  3362  (62);  Orient,  Long  Beach,  Latham  7425,  May  31,  1940 
(Latham);  Montauk  Point,  Latham  24172,  May  4,  1926  (Latham); 
Orient  Point,  Latham,  January  3,  1910,  (NYS):  Orient  Point,  Latham  8, 
January  9,  1911  (NYS). 

This  small-lobed  species  is  often  confused  with  X.  candelaria  (L.) 
Arn.  The  soredia  of  X.  fallax  are  produced  on  the  undersurface  as  well 
as  the  edges  of  more  or  less  hood-like  lobes,  whereas  the  soredia  or 
granules  of  X.  candelaria  are  produced  only  on  the  edges  of  the  lobes 
which  are  never  hood-like.  Thompson  (1949)  also  discusses  these 
differences. 

Xanthoria  fallax  is  a  rather  common  member  of  the  community 
on  roadside  elms.  It  was  found  once  on  concrete  (Brodo  2825)  at  Orient 
Point.  In  the  past,  this  species  must  have  been  common  in  the  New 
York  City  area  (see  citations  above). 

Distribution  —  Ontario  (leg.  LeBlanc),  Michigan  (seen  in  herb. 
MSC);  Wisconsin,  Oklahoma,  Arizona,  Black  Hills,  Saskatchewan,  Ca¬ 
nadian  archipelago:  Arctic-boreal  element;  Europe. 

232.  Xanthoria  parietina  ( L. )  Beltr.  Lich.  Bassan.  102.  1858.  Lichen 
parietinus  L.  Sp.  PI.  1143.  1753. 

Material  seen  —  KINGS  COUNTY:  Flatbush,  (Brainerd!)  1866 
(BKL).  SUFFOLK  COUNTY:  16  specimens  collected  by  Imshaug 
and/or  Brodo;  10  specimens  collected  by  Latham  (Latham);  Green- 
port,  Peck  (NYS);  Greenport,  Peck,  Sept.  (NYS);  Orient,  Young 
(BKL);  E.  Patchogue,  collector  unknown,  September  8,  1912  (Staten 
Island);  Sag  Harbor,  Britton  212,  July  17,  1898  (NY);  Sag  Harbor, 
Britton  211,  July  17,  1898  (NY). 


252  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

This  conspicuous  species,  although  usually  very  easily  identified, 
occasionally  shows  small-lobed,  richly  fruiting  forms  which  resemble 
X.  polycarpa  (Ehrh.)  Rieb.  The  latter  species,  however,  is  rarely  found 
along  the  coast  (never  on  Long  Island)  where  X.  parietina  is  most  com¬ 
mon.  In  addition,  X.  polycarpa  never  shows  any  tendency  towards  broad 
lobes  or  lack  of  apothecia,  and  typically  has  very  narrow,  finely-divided 
lobes  almost  obscured  by  apothecia.  Xanthoria  parietina  almost  always 
shows  some  broadened  lobes. 

Xanthoria  parietina  is  a  well  known  and  often  cited  example  of  a 
“nitrophilous”  or  “neutrophytic”  species,  and,  with  its  associated  species 
with  similar  requirements,  makes  up  the  well  known  Xanthorion  parie- 
tinae  alliance  discussed  in  full  by  Barkman  (1958)  and  also  by  des 
Abbayes  (1951).  Barkman  (1958)  does  not  consider  the  community 
halophytic,  despite  its  maritime  affinities,  although  he  notes  that  it  is 
“favored  by  salt  impregnation.”  He  prefers  to  call  it  “nitrophytic”  or 
“nitrophilous”  and  “subneutrophytic.”  The  observations  of  Maas  Ges- 
teranus  (1955)  of  X.  parietina  growing  well  on  the  windblown  edges  of 
salt  lakes  in  Kenya,  Africa,  might  point  to  the  importance  of  sodium, 
high  pH,  or  some  other  minerals  to  the  species. 

Both  Barkman  (1958)  and  Almborn  (1948)  note  that  the  com¬ 
munity  develops  best  on  the  road-facing  sides  of  trees  along  dusty  roads, 
especially  where  nitrogen-rich  dust  may  be  blown  on  the  substrata  and 
thallus,  although  Almborn  prefers  to  view  with  skepticism  the  theory 
that  nitrogen  concentration  is  the  chief  causal  factor  involved  in  this 
distribution.  On  Long  Island,  X.  parietina  was  also  most  commonly  found 
on  the  road-facing  side  of  roadside  trees.  Although  elms  seemed  to  be 
the  most  suitable  substrate,  it  was  also  collected  on  roadside  oaks  and 
maples. 

Des  Abbayes  (1934)  noted  X.  parietina  in  the  upper  hygrohaline 
(salt  spray)  zone  of  his  maritime  rock  community.  On  Long  Island,  its 
distribution  is  more  or  less  maritime,  often  being  found  very  close 
to  the  coast  (figure  80  and  p.  59). 

Distribution  —  On  northeastern,  Pacific  and  Gulf  coasts  (map: 
Hale,  1955c):  Temperate  element.  Oceanic  subelement(?)  Hale,  1961a); 
Europe,  in  lowlands  extending  far  inland  and  up  to  an  altitude  of  about 
1500m  (Maas  Gesteranus,  1955);  Asia  (Vainio,  1928,  Zahlbruckner, 
1930). 

59.  TELOSCHISTES  Norm. 

233.  Teloschistes  chrysophthalmus  (L.)  Beltr.  Lich.  Bassan.  109. 
1858.  Lichen  chrysophthalmos  (sic)  L.  Mantissa  PI.  2:  311.  1771. 

Material  seen  —  COUNTY  UNKNOWN:  Long  Island,  N.  Y„ 
Lloyd  (L.  1.  133)  (NY);  Long  Island,  G.  B.  Brainerd  (NYS).  SUF¬ 
FOLK  COUNTY:  Orient,  Latham,  May  20,  1914  (Latham);  Greenport, 
Peck  151,  Sept.  (NYS);  Greenport,  Peck  (NYS);  Moriches,  (Brainerd!) 
(BKL  032035);  Sayville,  Lloyd  (L.  I.  138),  December  3,  1896  (NY); 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  253 

Sag  Harbor,  Britton  210,  July  17,  1898  (NY);  Sag  Harbor,  on  Judge 
Daly’s  place,  Britton,  July  13,  1897  (NY). 

Teloschistes  chrysophthalmus  probably  was  a  member  of  the  coastal 
tree  community.  Along  with  T.  flavicans,  this  species  has  apparently  dis¬ 
appeared  from  Long  Island.  The  hurricane  of  1938  (see  Latham,  1945) 
was  probably  an  important  factor  in  cutting  down  the  population  size 
to  a  point  below  the  level  at  which  the  species  could  maintain  them¬ 
selves  without  reinvasion.  Since  Long  Island  appears  to  be  the  northern 
limits  for  both  species,  reinvasion  was  very  unlikely. 

Distribution  —  Widely  distributed  in  the  warm  areas  of  the  world 
(Zahlbruckner,  1931);  “Mexican  element,”  Texas  to  Minnesota  in  Great 
Plains  and  in  New  England  (Hale,  1961a):  Tropical  element,  Appa¬ 
lachian-Temperate  subelement;  Europe. 

234.  Teloschistes  flavicans  (Sw.)  Norm.  Nytt.  Mag.  Naturvid.  7:  229. 
1853.  Lichen  flavicans  Sw.  Nov.  Gen.  Sp.  PI.  147.  1788. 

Material  seen  —  SUFFOLK  COUNTY:  Orient  point,  Latham 
(CUP);  Orient  Point,  Latham  19,  April  9,  1910  (NYS). 

This  species  was  apparently  a  member  of  the  aerohaline  Juniperus 
community  (Latham,  pers.  comm.). 

Distribution  —  “Mexican  element,”  coastal  areas  of  Texas  and  the 
Carolinas  (Hale,  1961a);  Florida,  California  (Rudolph,  1955),  Con¬ 
necticut:  Tropical  element,  Coastal  Plain  subelement  (?);  Europe  (Poelt, 
1963);  Asia  (Zahlbruckner,  1930). 

PHYSCIACEAE 

60.  BUELLIA  De  Not. 

235.  Buellia  cu-tisii  (Tuck.)  Imsh.  in  Brodo,  comb.  nov.  Gyrosto- 
mum  curtisii  Tuck.  Amer.  J.  Arts  Sci.  II,  25:  430.  1858. 

Material  seen  —  SUFFOLK  COUNTY:  72  specimens  collected  by 
Imshaug  and/or  Brodo;  10  specimens  collected  by  Latham  (Latham). 

This  combination  was  first  used  in  a  thesis  by  Imshaug  (1951,  and 
1952  [abstract]).  Neither  usage,  however,  constituted  valid  publication. 
Culberson  (1953)  did  mention  the  new  combination,  but  since  he  did  not 
cite  the  basionym,  his  usage  also  does  not  constitute  valid  publication. 

This  species,  the  most  common  corticolous  Buellia  on  Long  Island, 
is  superficially  identical  with  B.  stillingiana.  The  two  species  have  very 
thin,  more  or  less  continuous,  greenish  grey  thalli  with  pitch  black 
apothecia.  Both  contain  norstictic  acid  and  both  are  found  on  a  variety 
of  deciduous  trees,  usually  smooth-barked  species,  and  usually  at  breast 
height.  The  differences  between  the  two  species  lie  in  their  apothecial 
and  spore  characters  (key,  p.  144). 

Distribution  —  Southeastern  United  States,  especially  along  the 
coastal  plain,  north  to  Connecticut  (map:  Imshaug,  1951):  Temperate 
element.  East  Temperate  subelement  (?);  endemic. 

236.  Buellia  dialyta  (Nyl.)  Tuck.  Gen.  Lich.  187.  1872.  Lecidea 
dialyta  Nyl.  Flora  52:  123.  1869. 


254  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1282  (21),  2460 

(22). 

Buellia  dialyta,  a  relatively  rare  species  on  Long  Island  and  else¬ 
where,  is  unusual  in  having  a  PD  +  red  thallus  reaction  (due  to  fumar- 
protocetraric  acid).  Its  thallus  is  usually  very  thin  and  scanty  (Imshaug, 
1951)  but  the  Long  Island  material  showed  fairly  well-developed  thalli, 
white  to  pale  ashy,  at  first  thin  but  becoming  thick  and  somewhat  rugose 
and  almost  granulose. 

The  species  was  found  once  on  the  bark  of  Quercus  velutina  and 
once  on  the  top  surface  of  a  rotting  log. 

Distribution  —  Maine,  Vermont,  New  Hampshire,  Massachusetts, 
Connecticut,  New  York,  Pennsylvania,  Tennessee,  California  (type  local¬ 
ity,  but  the  type  may  well  have  been  mislabelled)  (Imshaug,  1951): 
Temperate  element,  Appalachian  subelement,  Appalachian  unit;  endemic. 

237.  Buellia  polyspora  (Will,  in  Tuck.)  Vain.  Acta  Soc.  Faun.  FI. 
Fenn.  7(1):  171.  1890.  Buellia  myriocarpa  var.  polyspora  Will,  in  Tuck. 
Syn.  N.  Amer.  Lich.  2:  97.  1888. 

Material  seen  —  COUNTY  UNKNOWN:  Long  Island,  Latham, 
1914  (MSC).  SUFFOLK  COUNTY:  60  specimens  collected  by  Imshaug 
and/or  Brodo;  Orient,  Latham  20,  April  5,  1914  (Latham). 

Buellia  polyspora  differs  from  B.  punctata  sens.  str.  in  several  ways, 
in  addition  to  having  12  to  24  spores  per  ascus  rather  than  8.  The 
exciple  in  B.  polyspora  is  almost  hyaline  within  (as  in  B.  curtisii), 
whereas  in  B.  punctata,  the  exciple  is  solid  dark  brown  to  black.  In  addi¬ 
tion,  B.  polyspora  is  only  found  on  the  bark  of  deciduous  trees,  whereas 
B.  punctata  is  found  on  a  variety  of  substrates  including  lignum  and 
the  bark  of  conifers. 

On  Long  Island,  B.  polyspora  shows  a  limited  eastern  distribution, 
possibly  reflecting  extreme  intolerance  to  air  pollution,  but  more  likely 
indicating  its  preference  for  the  well  lighted,  open  woods  and  shrubby 
downs  most  common  in  that  part  of  the  island. 

Distribution  —  Throughout  eastern  United  States  (Imshaug,  1951): 
Temperate  element.  East  Temperate  subelement;  Brazil  ( Wainio,  1890). 

238.  Buellia  punctata  (Hoffm.)  Mass.  Ricerch.  Auton.  Lich.  81, 
f.  165.  1852.  Verrucaria  punctata  Hoffm.  Deutschl.  FI.  2:  192.  1796. 

Material  seen  —  COUNTY  UNKNOWN:  Long  Island,  N.  Y., 
Latham,  1914  (CUP).  SUFFOLK  COUNTY;  Imshaug  25582  (52), 
25616A  (116),  25770E  (121);  Orient,  Latham  62,  May  23,  1914 
(Latham);  Orient,  Latham  3941,  April  20,  1927  (Latham);  Orient,  Long 
Beach,  Latham  22333,  December  7,  1944  (Latham);  Orient  Point,  La¬ 
tham,  1911  (CUP). 

The  relationship  between  this  species  and  B.  polyspora  has  already 
been  discussed.  Buellia  punctata  is  very  rare  on  Long  Island,  being 
found  on  bark  (often  conifer  bark)  and  on  old  wood.  The  species  was 
also  collected  on  pine  bark  in  Cape  Cod. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


255 


Distribution  —  Throughout  United  States  and  southern  Canada,  and 
on  the  west  and  north  coasts  of  Alaska,  but  absent  from  all  other  parts 
of  high  boreal  and  arctic  Canada  (map:  Imshaug,  1951):  Temperate 
element  (?),  North  Temperate  subelement:  Europe;  Asia  (Zahlbruckner, 
1930). 

239.  Buellia  stigmaea  Tuck.  Syn.  N.  Amer.  Lich.  2:  90.  1888. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2672  (108).  2807 

( 106),  3079  (128),  3881  (62). 

This  species  is  very  similar  in  general  appearance  and  chemical 
reactions  to  Rhizocarpon  cinereovirens  which,  however,  has  a  greenish 
epithecium  and  apothecial  margin,  and  has  very  lightly  tinted,  slightly 
larger  spores,  each  usually  showing  a  conspicuous  gelatinous  sheath 
or  “halo.” 

Buellia  stigmaea  has  a  smooth,  whitish  grey  thallus  which  becomes 
irregularly  cracked  and  areolate,  allowing  a  conspicuous  black  prothallus 
to  show  through  between  the  areoles  as  well  as  beyond  the  thallus  edge. 
In  contrast,  Rhizocarpon  cinereovirens  has  a  dirty  grey-green  to  whitish 
grey  thallus  which  is  minutely  areolate  to  almost  granulose  and  lacks  a 
black  prothallus. 

Distribution  - —  Appalachians,  Missouri  (north  of  Ozarks)  (map: 
Imshaug,  1951);  Alaska  (Imshaug,  pers.  comm.)?:  Temperate  element, 
Appalachian  subelement,  Appalachian  unit  (?);  endemic. 

240.  Buellia  stillingiana  J.  Stein.  Oest.  Bot.  Zeitschr.  68:  144.  1919. 

Material  seen  —  NASSAU  COUNTY:  Cold  Spring,  Grout,  April  1, 

1900  (BKL).  SUFFOLK  COUNTY:  37  specimens  collected  by  Imshaug 
and/or  Brodo;  Greenport,  Latham,  May  12,  1960  (Latham);  Orient* 
Latham  736,  March  10,  1915  (Latham);  Orient,  Latham  737,  May  3* 
1915  (Latham);  Orient,  Latham  22331  A,  December  7,  1944  (Latham); 
Eastport,  Schrenk,  June  26,  1894  (MO). 

This  species,  long  confused  with  B.  parasema  (Ach.)  De  Not.,  is 
actually  quite  distinctive.  It  differs  from  B.  parasema  in  lacking  oil 
droplets  in  the  hymenium  and  in  having  a  “T”-shaped,  grey  apothecial 
stipe,  with  a  more  or  less  uniformly  brown-black  exciple  rather  than 
an  exciple  which  is  pale  brown  to  pale  olivaceous  within  (Imshaug,  1951; 
Lamb,  1954).  The  similarities  between  B.  stillingiana  and  B.  curtisii  are 
discussed  with  the  latter. 

Buellia  stillingiana  is  a  common  inhabitant  of  smooth-barked  de¬ 
ciduous  trees,  especially  in  well-lighted  situations. 

Distribution  —  Throughout  eastern  United  States,  and  in  Pacific 
Northwest  (map:  Imshaug,  1951);  absent  in  Black  Hills:  Temperate 
e'ement,  North  Temperate  subelement  (?);  endemic. 

241.  Buellia  turgescens  Tuck.  Gen.  Lich.  185.  1872. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1806  (127),  2371 
(123),  2673  (108),  3423  (134);  Orient,  Latham,  May  15,  1920  (La¬ 
tham);  Orient,  Long  Beach  factory  site,  Latham  22337,  December  7, 
1944  (Latham). 


256  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

This  distinctive  saxicolous  species  has  a  thick,  brownish  grey 
verrucose  to  granulose  thallus  with  very  small  apothecia  (less  than  0.5 
mm  in  diameter)  either  partially  sunken  into  the  verrucae  and  appearing 
aspicilioid,  or,  more  commonly,  sessile  and  prominent. 

It  grows  on  exposed  or  partially  shaded  granite  boulders,  and,  on 
one  occasion,  was  found  by  Latham  on  a  brick  (figure  65).  Lignicolous 
specimens  are  known  from  Massachusetts  (Imshaug,  1951). 

Distribution  —  Maine,  Massachusetts,  Connecticut,  New  York, 
Ohio,  Iowa,  Kansas,  Minnesota,  California,  Washington  (Imshaug, 
1951):  Temperate  element.  North  Temperate  subelement;  endemic. 

61.  RINODINA  (Ach.)  S.  Gray 

242.  Rinodina  applanata  Magn.  Bot.  Not.  43.  1947. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3077b  (128). 

This  species,  described  from  Louisiana,  was  found  only  once  on 

Long  Island  (on  oak  bark)  and  probably  is  rare  throughout  its  range. 
It  is  characterized  by  its  thin,  grey  thallus  and  uniformly  thick-walled 
spores  with  spherical  lumina  ( pachyspores;  figure  90c). 

Distribution  —  Louisiana  (type  locality),  Oklahoma:  endemic. 

243.  Rinodina  confragosa  (Ach.)  Korb.  Syst.  Lich.  Germ.  125.  1855. 
Parmelia  confragosa  Ach.  Meth.  Lich.  Suppl.  33.  1803. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2662b  (108),  3048 
(50). 

Rinodina  confragosa  is  a  rare  member  of  the  community  found  on 
shaded  boulders,  occurring  there  with  Rhizocarpon  intermedium  and 
Buellia  stigmaea.  Its  small  apothecia  (0. 5-1.0  mm  in  diameter)  are  flat  to 
slightly  concave.  The  apothecial  disks  are  brown  and  are  bounded  by 
a  prominent  rim  which  is  smooth  or  becoming  crenulate.  The  spores  are 
conspicuously  mischoblastiomorphic  (see  Imshaug,  1957c). 

Distribution  —  Massachusetts,  New  Jersey,  Louisiana,  Illinois,  Min¬ 
nesota,  Oregon,  California  (Fink,  1935);  Michigan,  Arizona,  Black 
Hills,  Washington,  Alaska:  Temperate  element.  North  Temperate  sub¬ 
element;  Europe;  Asia,  Africa  (Fink,  1910). 

244.  Rinodina  milliaria  Tuck.  Proc.  Arner.  Acad.  Arts  Sci.  12:  175. 
1877. 

Material  seen  —  SUFFOLK  COUNTY:  Imshaug  25615  (116), 
25627  (116),  25680  (72),  25687  (72),  25753b  (132);  Brodo  594  (92), 
1064  (130),  1792  (127),  2607  (84),  2726  (111),  2830  (115),  3108 
(122),  3317  (129);  Orient,  Long  Beach,  Latham  22,  April  16,  1914 
(Latham);  Orient,  Long  Beach,  Latham  22335,  December  7,  1944 

(Latham);  Orient,  Long  Beach,  Latham  22355aB,  December  1,  1944 

(Latham);  Orient,  Long  Beach,  Latham  22339B,  December  7,  1944 

(Latham);  Orient,  Brown's  Hills,  Latham  23057,  March  18,  1945  (La¬ 

tham);  Orient,  Latham  36804,  April  10,  1956  (Latham);  Southold, 
Latham  36952B,  October  10,  1960  (Latham). 

As  pointed  out  by  Magnusson  (1947),  the  apothecia  of  this  species 
often  lose  much  of  their  lecanorine  margins,  and  this,  together  with  the 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


257 


black  hypothecium,  often  give  specimens  the  appearance  of  a  Buellia.. 
However,  on  every  thallus  there  are  always  some  apothecia  showing  a 
typical  grey  or  greenish  margin. 

The  spores  are  normally  uniseptate,  8-15  x  5-7  p.,  but  one  specimen 
( Imshaug  25753b ),  showed  some  unusual  spores  mixed  in  with  the 
normal  ones.  These  aberrant  spores  were  3-septate  and  slightly  curved, 
and  measured  15-20  x  5-7  p.. 

Rinodina  milliaria  was  found  often  on  bark  and  wood,  especially 
in  windswept  and  salt-sprayed  areas.  Its  distribution  (figure  78)  reflects 
its  coastal  tendencies  on  Long  Island.  Although  it  appears  almost  mari¬ 
time  on  Long  Island,  the  species  has  been  reported  from  as  far  west  as 
Manitoba. 

Distribution  —  New  England,  New  York,  Pennsylvania  (Fink, 
1935);  Maine,  Wisconsin  (Magnusson,  1947);  Manitoba:  Temperate 
element.  North  Temperate  subelement;  endemic  (?). 

245.  Rinodina  oreina  (Ach.)  Mass.  Ricerch.  Auton.  Lich.  16,  f.  24. 
1852.  Lecanora  straminea  /}.  L.  oreina  Ach.  Lich.  Univ.  433.  1810. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  1801  (127),  1892 
(114),  2364  (123),  2659  (108),  2675  (108),  2734  (111),  2816  (106), 
3321  (129),  3447  (134),  3851  (76). 

All  the  Long  Island  material  of  this  species  showed  a  PD  — , 
C  +  red  thallus  reaction  and  thus  belongs  to  “Chemical  Strain  II”  of 
Hale  (1952b).  The  specimens  tested  microchemically  contained  gyro- 
phoric  acid. 

Rinodina  oreina  was  found  growing  on  exposed  granite  boulders 
in  eastern  Long  Island,  especially  near  the  bays  and  ocean. 

Distribution  —  Throughout  United  States,  boreal  and  arctic  Canada 
(map:  Hale,  1952b):  Arctic-boreal  element;  Europe.  The  gyrophoric 
acid  strain,  by  itself,  has  a  North  Temperate  distribution  (see  Hale, 
1952b). 

246.  Rinodina  pachysperma  Magn.  Bot.  Not.  193.  1953. 

Material  seen  —  SUFFOLK  COUNTY:  16  specimens  collected  by 

Imshaug  and/or  Brodo;  Orient,  North  Locust,  Latham  17,  April  6, 
1914  (Latham);  Shelter  Island,  collector  unknown,  October  10,  1910 
(FH). 

The  Long  Island  material  was  compared  with  the  type  specimen 
from  Wisconsin  (in  herb.  J.  Thomson).  The  spore  and  apothecial 
characters  agreed,  but  most  of  the  eastern  specimens  had  a  smoother 
thallus,  frequently  in  flat  areoles  which  sometimes  partially  lift  off  the 
surface  and  appear  subsquamulose.  The  type  specimen  has  a  well-de¬ 
veloped,  minutely  areolate  thallus,  but  showed  the  same  olive-green  to 
dark  green  color.  Tendencies  toward  intergradation  were  seen,  however, 
and  no  really  significant  differences  could  be  found. 

There  are  a  few  characters  which  appeared  to  be  constant  and 
conspicuous  which  deserve  special  mention.  The  apothecia  often  show 
both  proper  and  thalline  margins  in  macroscopic  view.  The  spores  were 


258  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

variable  in  shape,  often  slightly  curved,  rounded  at  one  end  and  pointed 
at  the  other,  or  merely  ellipsoid. 

The  species  was  collected  on  the  bark  of  various  deciduous  trees, 
usually  close  to  the  coast  (figure  79). 

Distribution  —  Wisconsin  (type  locality),  Black  Hills;  endemic. 

247.  Rinodina  salina  Degel.  Uppsala  Univ.  Arsskr.  192.  1939  (nom. 
nud.);  Ark.  Bot.  30A  (1):  55.  1940. 

Material  seen  —  SUFFOLK  COUNTY;  Brodo  2828 A  (115); 
Orient,  Latham  2873  (22264),  February  7,  1926  (Latham)  (with  Calo- 
placa  fiavovirescens) . 

Degelius  (1939)  pointed  out  that  the  name  Rinodina  demissa  Arn., 
under  which  this  plant  has  generally  been  considered,  cannot  be  used 
since  the  basionym.  Zeora  metaholica  demissa  Fldrke,  is  based  on  an 
entirely  different  taxon,  Bnellia  ambigua  (Ach.)  Malme. 

Rinodina  salina,  a  well-known  maritime  lichen  in  Europe,  was  first 
reported  from  North  America  by  Riisanen  (  1933)  (as  R.  demissa ),  from 
New  Brunswick,  Canada,  and  then  by  Degelius  (1940)  from  Prince’s 
Point  in  Maine,  where  it  was  found  "on  maritime  rocks  in  the  middle- 
hygrohaline.”  The  original  description  notes  the  thallus  as  brownish  to 
dark  ashy.  A  specimen  seen  at  the  U.S.  National  Museum  showed  a 
distinct  brownish  tint.  The  Long  Island  specimens  had  no  brown  tint 
at  all;  they  were  whitish  to  ashy,  becoming  sordid  ashy. 

As  in  Europe  and  Maine,  the  Long  Island  specimens  were  in  the 
maritime  zone,  specifically,  the  aerohaline  zone.  Both  specimens  were 
on  calcareous  substrates  (concrete  and  bone)  and  were  associated  with 
species  of  Caloplaca. 

Distribution  - —  New  Brunswick,  Maine  (see  above).  Black  Hills 
(a  very  odd  record  in  view  of  its  ecology);  Europe. 

62.  PYXINE  Fr. 

248.  Pyxine  sorediata  (Ach.)  Mont,  in  Sagra,  Hist.  Cuba  8:  188. 
pi.  7,  f.  4.  1838-42.  Lecidea  sorediata  Ach.  Syn.  Lich.  54.  1814. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-255  (67),  869 
(47),  1886  (114),  2056  (45),  2494  (67),  3895  (112). 

Imshaug  (1957c)  has  thoroughly  discussed  this  species  as  it  occurs 
in  North  America.  None  of  the  Long  Island  material  had  apothecia. 
Pyxine  sorediata  was  found  mostly  at  breast  height  on  oaks  in  pine-oak 
forests. 

Distribution  —  Throughout  eastern  United  States  (map:  Imshaug, 
1957c):  Temperate  element.  East  Temperate  subelement;  eastern  Asia 
( ibid ) . 

63.  PHYSCIA  (Schreb.)  DC. 

249.  Physcia  adscendens  (Th.  Fr.)  Oliv.  FI.  Lich.  Orne  1:  79.  1882. 
Physcia  stellaris  var.  adscendens  Th.  Fr.  Lich.  Scand.  1  :  138.  1871. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  59-330  (53),  2117 
(93),  2120  (NE.  of  71);  Orient  Point,  Latham  30,  April  18,  1910 
(NYS). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


259 


This  species  was  listed  and  discussed  by  Fink  (1935)  under  Physcia 
hispida  (Schreb.)  Frege,  a  name  which  cannot  be  used  due  to  its  uncer¬ 
tain  meaning  (Thomson,  1963). 

Physcia  adscendens  is  relatively  rare  on  Long  Island.  As  is  usual 
for  the  species,  it  occurs  as  a  member  of  the  communities  on  roadside 
poplars  and  elms  or  calcareous  rocks  (the  Xanthorion  parietinae  alli¬ 
ance  as  recognized  in  Europe). 

Distribution  —  Throughout  northern  and  western  United  States  and 
southern  Canada  (map:  Thomson,  1963):  Temperate  element.  North 
Temperate  subelement,  Europe,  Asia  (ibid). 

250.  Physcia  aipolia  (Ehrh.  in  Humb.)  Hampe  in  Furnr.  Naturh. 
Topogr.  Regensburg  2:  249.  1839.  Lichen  aipolius  Ehrh.  in  Humb.  FI. 
Friburg.  Spec.  19.  1793. 

Material  seen  —  SUFFOLK  COUNTY:  21  specimens  collected  by 
Imshaug  and/or  Brodo. 

The  white  spots  characteristic  of  this  species  can  best  be  termed 
maculae,  rather  than  pseudocyphellae  as  they  are  sometimes  called. 
In  the  latter,  the  upper  cortex  must  be  broken,  allowing  medullary 
hyphae  to  reach  the  surface.  This  is  not  the  case  in  P.  aipolia,  where  the 
spots  appear  to  be  tiny  discontinuities  in  the  algal  layer  beneath  a  con¬ 
tinuous  upper  cortex. 

Physcia  aipolia  is  a  member  of  the  community  on  well-lighted 
black  oaks  and  is  usually  found  in  open  pine-oak  forests  (figure  46). 

Distribution  —  Throughout  the  United  States,  southern  Canada, 
and  coastal  Alaska  (map:  Thomson,  1963):  Temperate  element.  North 
Temperate  subelement,  Europe,  Asia  (ibid). 

251.  Physcia  millegrana  Degel.  Ark.  Bot.  30A(  1 ) :  56.  1940. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  {Brainerd  ?)  (BKL 

032039).  NASSAU  COUNTY:  Brodo  1307  (15),  3195  (7).  SUFFOLK 
COUNTY:  64  specimens  collected  by  Imshaug  and/or  Brodo;  Green- 
port,  Latham,  June  26,  1960  (Latham);  Orient,  Latham  68,  May  23, 
1914  (Latham);  Orient,  Latham  7453,  June  5.  1933  (Latham);  Orient, 
Long  Beach,  Latham  8586,  April  25,  1939  (Latham);  Greenport,  Latham 
36928,  June  26,  1960  (Latham);  Orient,  Latham  36936,  September  10, 
1960  (Latham). 

This  very  common  species  was  for  years  considered  to  be  the  same 
as  the  European  P.  tribacia  (Ach.)  Nyl.  (see  Fink,  1935).  The  two  are 
separated  on  the  basis  of  their  lower  cortices:  paraplectenchymatous  in 
P.  millegrana,  and  not  paraplectenchymatous  in  P.  tribacia.  The  only 
North  American  record  of  P.  tribacia  is  from  the  Northwest  Territories 
(Thomson,  1963). 

The  thallus  varies  from  having  flat,  very  finely-dissected  lobes  with 
very  sparse  marginal  granules  to  a  form  having  densely  granular  lobe 
margins,  often  piling  up  in  the  thallus  center  and  almost  giving  the 
appearance  of  a  granular  crust. 


260  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  species  usually  is  found  on  the  bark  of  a  variety  of  deciduous 
trees,  especially  in  well-lighted  forests  (figure  71).  It  was  found  occa¬ 
sionally  on  the  bark  of  Juniperus  virginiana  (which  often  supports 
neutrophytic  communities),  and  once  on  a  granite  boulder  (Brodo 
3096).  When  the  species  grows  on  rock,  it  can  be  confused  with  P.  sub- 
filis.  The  separation  of  the  two  is  discussed  with  the  latter  species. 

Distribution  —  Eastern  United  States  south  to  North  Carolina 
and  Texas,  California  (introduced)  (map:  Thomson,  1963):  Temperate 
element,  East  Temperate  subelement,  endemic  (ibid). 

252.  Physcia  orbicularis  (Neck.)  Potsch  in  Potsch  &  Schiederm. 
Syst.  Aufzahl.  Samenlos.  Pfl.  247.  1872.  Lichen  orbicularis  Neck.  Meth. 
Muse.  88.  1771. 

Material  seen  —  NASSAU  COUNTY:  Brodo  1313  (15),  1501 
(9),  1502  (14).  SUFFOLK  COUNTY:  52  specimens  collected  by 
Imshaug  and/or  Brodo:  Orient,  Long  Beach,  Latham ,  April  16,  1933 
(Latham);  Orient,  Latham,  April  15,  1914  (Latham);  Orient,  Latham 
892,  February  1,  1920  (Latham);  Manorville,  Latham  8622,  May  20, 
193(8)?  ( Latham );  Montauk,  Latham  31912,  January  5,  1953  (Latham); 
Orient  Point,  Latham  8,  January  9,  1911  (NYS). 

There  are  two  common  forms  of  P.  orbicularis  in  North  America, 
one  with  a  white  medulla  (f.  orbicularis),  and  one  with  on  orange-red 
medulla  ( f.  rubropulchra  Degel.).  Both  forms  are  represented  on  Long 
Island,  with  the  latter  being  much  more  abundant.  Forma  rubropulchra 
appears  to  be  restricted  to  North  America  but  corresponds  to  f.  Hueana 
(Harm.)  Erichs,  in  Europe,  which  has  a  yellow-orange  medulla  with 
soredia  of  the  same  color  (Degelius,  1940).  A  critical  study  of  the 
anthraquinone  pigments  involved  in  these  vicarious  forms  might  prove 
very  interesting  in  light  of  some  of  the  recent  data  regarding  the  system¬ 
atic  and  biogenetic  importance  of  pairs  of  closely  “related”  depsides 
and  depsidones  in  closely  related  taxa  (see  Runemark,  1956;  C.  Cul¬ 
berson  1963,  1964;  Imshaug  and  Brodo,  1966). 

Bruce  Fink  (1935)  apparently  referred  to  f.  orbicularis  as  P.  virella 
(Ach.)  Flag.,  and  to  f.  rubropulchra  as  “P.  endochrysea  (Hampe)  Nyl.” 
( P .  endochrysea  (Nyl.)  Hampe  in  Krempelh.) .  His  P.  endochrysea  may 
have  also  included  some  P.  endococcinea  (Korb.)  Th.  Fr.  (a  saxicolous 
non-sorediate  species),  since  Fink  lists  P.  obscura  f.  endococcinea  as  a 
synonym  of  P.  endochrysea  and  Zahlbruckner  (1931)  lists  the  former 
as  a  synonym  of  P.  endococcinea. 

Tuckerman  (1882)  regarded  P.  obscura  var.  endochrysea  Nyl.  as 
synonymous  with  his  P.  obscura  var.  erythrocardia  Tuck.  Thomson 
(1963),  who  I  assume  saw  Tuckerman’s  type,  refers  var.  erythrocardia  to 
P.  ciliata  (as  f.  erythrocardia  [Tuck.]  Thoms.).  Degelius  (1941)  regarded 
“P.  endochrysea  Krempelh.”  as  questionably  synonymous  with  P.  ciliata 
var.  erythrocardia.  Hale  and  Culberson  (1960)  listed  P.  endochrysea  as 
a  synonym  of  P.  orbicularis  f.  rubropulchra. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


261 


Thus,  the  epithet  endochrysea  has  been  used  for  at  least  three  North 
American  species  which  have  forms  with  a  red  medulla  (P.  orbicularis, 
P.  endococcinea,  and  P.  ciliata).  Although  the  epithet  endochrysea  is  of 
no  nomenclatural  importance  as  far  as  these  species  are  concerned,  it 
would  be  well  for  its  identity  to  be  established.  Thomson  (1963)  did  not 
mention  the  epithet  at  all,  and  so  a  final  solution  to  the  problem  must 
await  an  examination  of  the  type. 

Physcia  orbicularis  is  found  most  often  on  shaded  tree  bases,  but  is 
also  found  at  other  vertical  positions  on  a  variety  of  trees.  It  has  also  been 
collected  on  cement  foundations. 

Distribution  —  Throughout  United  States  and  adjacent  Canada, 
especially  in  east  (map:  Thomson,  1963):  Temperate  element,  North 
Temperate  subelement,  Europe,  Asia  (ibid).  Forma  orbicularis,  as  in 
species;  forma  rubropulchra :  East  Temperate,  endemic  (ibid). 

253.  Physcia  stellaris  (L.)  Nyl.  Act.  Soc.  Linn.  Bordeaux  21:  307. 
1856.  Lichen  stellaris  L.  Sp.  PI.  1144.  1753. 

Material  seen  —  SUFFOLK  COUNTY:  26  specimens  collected  by 
Imshaug  and/or  Brodo;  Orient,  Long  Beach,  Latham  27219,  May  15, 
1947  (Latham);  Orient,  Long  Beach,  Latham  22330,  December  7,  1944 
(Latham);  Orient  Point,  Latham  26,  April  18,  1910  (NYS);  Orient 
Point,  Latham,  May  2,  1910  (NYS);  Greenport,  Peck  (NYS);  Sag 
Harbor,  Britton  211,  July  17,  1898  (NY). 

'Phis  species  was  found  on  the  bark  of  several  species  of  deciduous 
trees,  mainly  in  welLlighted  forests  (figure  47).  It  also  occurs  on  the 
bark  of  Juniperus  virginiana. 

Distribution  —  Throughout  United  States,  southern  Canada,  coastal 
Alaska,  central  Mexico  (map:  Thomson,  1963):  Temperate  element, 
North  Temperate  subelement,  Europe,  Asia  (ibid). 

254.  Physcia  subtilis  Degel.  Ark.  Bot.  30A(3):  72.  1941. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  2370  (123),  2654 

(108),  3356  (62),  3431  (134). 

Physcia  subtilis,  a  saxicolous  species,  is  similar  in  some  respects  to 
P.  millegrana,  although  the  latter  rarely  occurs  on  rock.  Some  of  their 
differences  are  outlined  below: 


Lobe  width 
Soredia  (granular) 
KOH  reaction 
of  medulla 
Anatomy 


Physcia  subtilis 
0.1 -0.5  mm 
marginal  and  apical 
+  (yellow) 

paraplectenchymatous 

throughout 


Physcia  millegrana 
0.3-1. 0(-1.5)mm 
only  marginal 


medulla  not 
paraplectenchymatous 


Degelius  (1941)  and  Thomson  (1963)  give  the  lobe  size  as  no 
broader  than  0.2  mm,  but  the  Long  Island  material  becomes  at  least 
twice  that  broad  in  a  few  cases. 


262  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Physcia  subtilis  was  found  on  granitic  rocks. 

Distribution  —  Eastern  United  States,  single  localities  in  Arizona 
and  Washington  (map:  Thomson,  1963):  Temperate  dement,  East 
Temperate  subelement  (?);  endemic. 

255.  Physcia  tribacoides  Nyl.  Flora  52:  322.  1869. 

Material  seen  —  SUFFOLK  COUNTY :  Imshaug  25578  (52) ; 
Brodo  2498  (67),  3904  (112);  Orient,  Latham  949B,  May  10,  1923 
(Latham). 

This  species  was  relatively  rare  on  Long  Island,  where  it  was  found 
on  bases  of  Quercus  velutina  and  Q.  alba  in  oak  forests  and  once  on 
Juniperus. 

Distribution  —  Eastern  United  States,  and  a  single  locality  in  Cali¬ 
fornia  (map:  Thomson,  1963):  Temperate  element,  East  Temperate  sub- 
element;  rare  in  Europe  (ibid). 

64.  ANAPTYCHIA  Korb. 

256.  Anaptychia  obscurata  (Nyl.)  Vain.  Acta  Soc.  Faun.  FI.  Fenn. 
7  (1):  137.  1890.  Physcia  obscurata  Nyl.  Ann.  Sci.  Nat.  Bot.  IV,  19: 
310.  1863. 

Material  seen  —  SUFFOLK  COUNTY:  Brodo  3908  (112);  Orient, 
Latham,  March  21,  1915  (Latham);  Orient,  Latham  949 ,  May  10,  1923 
( Latham) . 

This  species  has  been  treated  in  some  North  American  literature  as 
A.  heterochroa  Vain,  and  A.  sorediifera  ( Mull.  Arg.)  Du  Rietz  and 
Lynge  in  Lynge.  Kurokawa  (1962)  discusses  the  nomenclature  and 
morphology  of  A.  obscurata  in  great  detail.  In  addition,  Degelius  (1941) 
presents  a  thorough  treatment  of  the  separation  of  this  species  (sub 
A.  sorediifera)  from  the  often  confusing  A.  pseudospeciosa  (sub  A. 
speciosa ) . 

The  yellow  anthraquinone  pigment  on  the  lower  surface  of  A. 
obscurata  varies  in  concentration  from  one  part  of  the  species’  range 
to  another,  but  all  three  Long  Island  specimens  show  a  distinct  dark 
yellow  color  which  was  clearly  KOH  +  red-purple. 

The  species  is  rare  on  Long  Island  and  is  found  on  mossy  tree  bases. 

Distribution  —  "...  widely  distributed  in  tropical  and  temperate 
zones  around  the  world,”  eastern  North  America  (Kurokawa,  1962): 
Tropical  element,  Appalachian-Temperate  subelement;  Europe,  Asia, 
Africa  (ibid). 

257.  Anaptychia  paimulata  ( Michx.)  Vain.  Termeszetr.  Fiizeteck  22: 
299.  1899.  Psoroma  paimulata  Michx.  FI.  Bor.  Amer.  2:  321.  1803. 

Material  seen  —  KINGS  COUNTY:  New  Lots,  ( Brainerd ?)  (BKL 
032038). 

Fink  (1935)  probably  treated  this  species  under  the  name  A.  aquila 
(Ach.)  Mass.  The  latter  is  a  synonym  of  A.  fusca  (Huds.)  Vain.,  a 
European  species  (Kurokawa,  1962).  Anaptychia  paimulata  is  discussed 
in  detail  by  Kurokawa  (1962). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


263 


Distribution  —  Appalachian  Mountains  and  Great  Lakes  region 
(map:  Hale,  1956c,  sub.  A.  palmatula ):  Temperate  element,  Appalachian 
subelement,  Appalachian-Great  Lakes  unit;  Asia  (Kurokawa,  1962). 

258.  Anaptychia  pseudospeciosa  Kurok.  J.  Jap.  Bot.  34:  176.  1959. 

Material  seen  —  SUFFOLK  COUNTY;  Brodo  1390  (65). 

This  species  was  segregated  from  A.  speciosa  (Wulf. )  Mass,  by 
Kurokawa  (1959,  1962)  on  the  basis  of  the  former's  smaller  spores  and 
sorediate  apothecial  margin.  (A.  speciosa  has  spores  over  30^  long  and 
has  a  smooth,  crenulate,  apothecial  margin.)  Sterile  specimens  of  the 
inactive  form  (without  salacinic  acid)  are  virtually  identical  to  European 
A.  speciosa ,  and  there  is  growing  doubt  that  the  two  are  distinct.  The 
Long  Island  specimen  was  without  salacinic  acid.  It  was  found  on  Quer- 
cus  alha  close  to  the  base. 

Distribution  —  Tropical  and  subtropical  areas  throughout  the  world 
(Kurokawa,  1962),  eastern  United  States  (Kurokawa,  1959):  Tropical 
element,  Appalachian-Temperate  subelement;  Africa,  Asia  (Kurokawa, 
1962). 

65.  LEPRARIA  Ach. 

259.  Lepraria  incana  (L.)  Ach.  Lich.  Suec.  Prodr.  7.  1798.  Byssus 
incana  L.  Sp.  PI.  1169.  1753. 

Material  seen  —  NASSAU  COUNTY:  Brodo  551  (12).  SUFFOLK 
COUNTY:  Brodo  964  (S.  of  50),  3826  (66). 

Laundon  (1962)  gave  a  short  discussion  on  the  correct  status  of  the 
genera  Crocynia  and  Lepraria  and  noted  that  Crocynia  aeruginosa  Hue 
is  a  synonym  of  L.  incana. 

The  presence  of  easily  identifiable  depsides  and  depsidones  in  the 
Lepraria  complex  promises  to  help  clear  up  some  of  the  taxonomic  prob¬ 
lems  in  the  group.  Although  the  three  species  of  Lepraria  on  Long  Island 
can  be  identified  by  their  gross  morphology  alone,  they  can  also  be 
separated  by  their  chemical  constituents.  By  chromatographic  analysis 
all  specimens  of  L.  incana  were  found  to  contain  atranorin,  and  one 
from  southern  New  Jersey  ( Brodo  3546)  also  contained  fumarprotoce- 
traric  acid.  The  chemistry  of  the  latter  specimen  was  carefully  rechecked 
to  avoid  the  possibility  of  a  contaminant. 

Lepraria  incana  was  found  on  tree  bark  in  shaded  habitats. 

Distribution  —  New  Jersey,  but  probably  is  more  widely  distributed. 

260.  Lepraria  zonata  sp.  nov.  Crocynia  zonata  Near.  Lichen  Book 
354.  1947  (nom.  nud.). 

Material  seen  —  NASSAU  COUNTY:  Brodo  3514  (10).  SUF¬ 
FOLK  COUNTY:  Imshaug  25601  (SW  of  106);  Brodo  3844  (76), 
3869  (62)  (HOLOTYPE),  3876  (62). 

This  species  was  first  mentioned  in  the  literature  by  Nearing  (1947), 
who  presents  a  thorough  description  in  English  and  states  that  the  name 
is  “current  in  the  New  York  area”  but  has  an  "obscure  origin.”  Since  it 
appears  to  be  a  good  species  easily  recognized  by  morphology  and  chem¬ 
istry,  a  Latin  description  is  provided  here  to  make  the  name  valid. 


264  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Thallus  crustaceus,  omnis  granulatus  sed  conspicuus  segre- 
gatus  et  marginibus  lobatusculis,  series  irregulariter  areolatus; 
granulae  50-100  [t.  diain.,  sine  ascocarpi.  Materiae  chemicae:  acidum 
fumarprotocetraricum  et  atranorin.  Saxicola. 

Thallus  crustaceous,  entirely  granular,  but  clearly  delimited  and 
somewhat  lobed  at  edges,  becoming  irregularly  areolate;  granules  50  to 
IOOij.  in  diameter;  without  ascocarps.  Thallus  contains  fumarprotoce- 
traric  acid  and  atranorin  (by  chromatographic  analysis). 

Holotype:  NEW  YORK,  Suffolk  County:  Wading  River:  Brodo 
3869,  August  11,  1962,  vertical  surface  of  partially  shaded  boulder 
(MSC)  (figure  86). 


Figure  86.  Lepraria  zonata  (holotype).  Scale  equals  1  mm.  Drawing 
by  Brenda  Carter  Haas. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


265 


Some  specimens  show  the  delimited,  lobed,  thallus  edges  more 
clearly  than  others,  but  none  are  entirely  effuse.  The  Long  Island  speci¬ 
mens  all  contained  fumarprotocetraric  acid  and  atranorin  except  one 
(Brodo  3514),  which  showed  only  barbatolic  acid. 

This  species  is  found  on  partially  shaded  granite  boulders. 

Distribution  —  Northeastern  states  (Nearing,  1947),  Massachusetts 
(Cape  Cod)  (Brodo  4162). 

261.  Lepraria  sp. 

Material  seen  —  NASSAU  COUNTY:  Brodo  3499  (4).  SUFFOLK 
COUNTY:  Brodo  591  (92),  2620  (84). 

This  species  is  often  found  in  herbaria  under  Crocynia  membranacea 
(Dicks.)  Zahlbr.  and  apparently  is  quite  common.  The  Long  Island  spe¬ 
cies  differs  from  true  Lepraria  membranacea  in  forming  thick,  leprose 
mats,  never  membranous,  subsquamulose  sheets.  The  two  apparently 
differ  also  in  chemistry,  although  too  few  reliable  specimens  of  L.  mem¬ 
branacea  have  been  examined  to  permit  generalization. 

Culberson  (1963a)  reported  a  leprose,  pale  green  crust  growing  on 
greenhouse  pots  in  Paris  which  produced  stictic  acid  and  atranorin.  There 
is  a  very  good  chance  that  this  as  yet  unidentified  species  from  the 
United  States  may  be  the  same  as  the  one  from  Europe.  There  is  obvi¬ 
ously  a  great  need  for  a  thorough  investigation  of  the  North  American 
Leprariae,  especially  one  done  with  a  careful  eye  for  thallus  chemistry. 

On  Long  Island,  the  species  is  found  on  moist,  shaded  tree  bases. 

Distribution  —  Unclear,  but  probably  widely  distributed. 


General  Discussions 


DISTRIBUTION  OF  LICHENS  ON  LONG  ISLAND 

In  summarizing  Long  Island  distributions  of  the  various  lichens,  a 
number  of  patterns  were  seen  to  recur.  The  patterns  correlated  with 
various  factors,  among  them  substrate  (tree,  rock,  or  soil)  distribution, 
vegetation  types,  climatic  gradients  such  as  the  fog  belt,  the  moraines, 
the  coastal  maritime  zones,  and  city  effects. 

1.  Substrate.  Substrate  distribution  will  define  a  species’  distribution 
limits  according  to  the  substrate  specificity  of  that  species.  A  substrate 
specific  species  may  be  secondarily  limited  by  climatic  or  other  factors 
within  the  range  of  the  substrate,  but  cannot  occur  outside  the  substrate 
range.  Thus,  a  species  completely  or  strongly  confined  to  pine  will  not 
extend  beyond  the  limits  of  the  distribution  of  pine.  Such  is  the  case 
with  Parmeliopsis  placorodia,  Lecidea  scalaris,  and  L.  anthracophila,  all 
of  whose  distributions  are  limited  to  the  pine  forests  which  in  turn 
reflect  the  natural  distribution  of  Pinus  rigida  on  Long  Island  (figures  45, 
43,  42).  There  are  several  species  closely  associated  with  oaks,  but  they 
are  limited  by  light  and/or  moisture  conditions  and  so  do  not  follow  the 
full  oak  distribution.  Graphis  scripta,  for  example,  is  only  found  in  the 
red  oak  forest  where  the  shade  and  humidity  is  greater  than  in  the  pine- 
oak  forest  ( Brodo,  1961a)  (figure  61).  The  opposite  is  true  of  Parmelia 
perforata.  P.  galbina,  Physcia  stellaris,  Ph.  aipolia.  Ph.  millegrana  (figures 
52,  51,  47,  46,  71).  Culberson  (1955b),  while  pointing  out  that  certain 
species  in  the  Wisconsin  lichen  flora  (e.g.,  Candelaria  concolor,  and 
Parmelia  aurulenta )  are  apparently  restricted  in  their  distribution  by  their 
substrate  preferences,  maintains  that  most  species  showing  clear-cut  dif¬ 
ferences  in  north-south  distribution  in  Wisconsin  are  responding  to 
climatic  differences  and  not  substrate  distributions. 

Certain  species  confined  to  bog  habitats  are  only  found  on  Chamae- 
cyparis  thyoides  and  Vaccinium  corymbosum.  Whether  these  species 
(such  as  Cetraria  ciliaris,  C.  viridis,  and  Usnea  trichodea )  are  restricted 
to  the  bogs  because  these  substrates  are  only  found  there,  or  whether  the 
climate  of  the  bog  is  the  determining  factor  is  still  unclear,  but  the  prob¬ 
lem  might  be  solved  at  least  partially  by  a  tran-plant  experiment  such  as 
was  described  on  p.  20. 

Certain  obligatory  saxicolous  species  have  distributions  reflecting  the 
positions  of  the  two  terminal  moraines  which  laid  down  large  numbers 
of  granitic  boulders  (erratics)  and  large  pebbles  and  stones  (p.  6). 

There  are  three  major  types  of  parent  soil  material  represented  on 
Long  Island:  hilly  glacial  till,  sandy  loam,  and  sand.  Over  these  there 
have  been  local  accumulations  of  alluvium,  beach  sands,  and  organic 
matter.  The  three  parent  materials  are  distinct  enough  to  influence  the 
distributions  of  several  terricolous  species,  and  yet  have  enough  char¬ 
acteristics  in  common  (e.g.,  sandy  texture,  excessive  drainage,  and  low 


267 


268 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


pH)  to  permit  certain  lichens  with  broader  ecological  limits  to  inhabit 
almost  any  part  of  the  island  having  exposed  glacial  parent  material. 

Baeomyces  roseus,  Cladonia  squamosa,  C.  caespiticia,  and  C.  pleu- 
rota  (figures  53,  54,  55,  56)  are  mainly  confined  to  the  hilly  glacial  till 
on  which  the  Plymouth-Haven  soil  association  has  been  formed,  with 
occasional  occurrences  on  the  sandy  loams  of  the  Bridgehampton  asso¬ 
ciation.  Cladonia  strepsilis  and  Pycnothelia  papillaria  (figures  72,  73)  are 
found  throughout  the  island  on  soils  of  various  types  exposed  by  natural 
or  man-made  erosion.  Cladonia  boryi,  C.  submitis,  and  C.  uncialis  are 
fairly  restricted  to  the  Colton  and  Adams  sandy  soil  association  (figures 
74,  75,  76). 

This  sand  dune  —  sand  plain  community,  although  abundant  on  the 
dunes  of  the  south  shore,  is  entirely  absent  from  those  on  the  north 
shore,  even  though  the  sand  texture  is  the  same  and  many  of  the  associ¬ 
ated  vascular  dune  plants  are  the  same.  The  reason  may  relate  to  the 
relative  salt  spray  deposition  in  the  two  areas  (p.  203). 

2.  Climate,  a.  Temperature:  The  differences  in  temperature  over 
Long  Island,  as  indicated  by  published  synoptic  temperature  maps,  are  too 
slight  to  influence  lichen  distribution  directly.  The  indirect  effects,  e.g., 
with  regard  to  moisture,  are,  however,  of  considerable  importance. 

The  warm  temperatures  of  the  city,  in  combination  with  its  relatively 
severe  fluctuations  in  humidity,  possibly  contribute  to  the  poor  lichen 
flora  in  western  Long  Island.  Rydzak  (1958)  believes  drought  to  he  the 
main  or  entire  cause  of  “lichen  deserts”  in  towns. 

The  relatively  cool  summer  temperatures  and  mild  winter  tempera¬ 
tures  of  eastern  Long  Island,  in  combination  with  the  very  high  humidity 
of  that  region,  contribute  towards  the  creation  of  a  distinctly  “oceanic” 
aspect  to  the  lichen  vegetation  in  that  area.  Faegri  (1958)  stressed  the 
importance  of  considering  both  temperature  and  relative  humidity  in  an 
area  suspected  of  being  oceanic  (see  discussion  below  under  “humidity”). 
Bog  habitats  also  have  this  combination  of  cool  temperatures  and  high 
humidity,  and  thus  provide  conditions  suitable  for  the  establishment  of 
oceanic  and  northern  species  west  of  the  highly  humid  cool  areas  in  the 
Montauk  region  (figure  31). 

b.  Precipitation:  Precipitation  differences  in  eastern  and  western 
Long  Island  are  also  small,  and  it  is  unlikely  that  rainfall  or  snowfall 
has  an  appreciable  direct  effect  on  lichen  distributions  on  Long  Island  as 
a  who'e.  The  secondary  effects  of  rainfall,  e.g.,  in  raising  the  air  humidity 
in  certain  forest  types  more  than  in  others,  can  and  probably  does  have 
an  effect  in  some  distributions. 

c.  Humidity:  Previous  workers  who  have  studied  coastal  lichen  vege¬ 
tation  (Barkman,  1958;  Almborn,  1948;  Degelius,  1935;  Mitchell,  1961) 
have  all  pointed  out  the  importance  of  air  humidity  on  the  distribution  of 
lichens.  They  particularly  cite  fog  frequency  as  being  an  important  factor. 
Degelius  (1935)  considers  hygric  factors  as  the  most  important  in  deter¬ 
mining  the  distribution  of  oceanic  species. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


269 


A  map  showing  the  numbers  of  days  on  Long  Island  with  dense  fog 
(figure  5)  strikingly  reflects  the  distribution  of  many  lichens  (figures 
64-70).  The  south  fluke  (fork)  of  Long  Island  has  a  much  greater 
species  richness  than  any  part  of  the  north  fluke  except,  perhaps,  for 
intensely  collected  Orient  Point.  Such  oceanic  species  as  Leptogium  cya- 
nescens,  Nephroma  laevigatum,  Loharia  qitercizans,  and  L.  pulmonaria 
all  show  distributions  more  or  less  restricted  to  the  fog  belt  (figures  5 
and  31 ). 

d.  Salt  Spray:  The  maritime  coastal  distributions  illustrated  by  Calo- 
placa  citrina,  Rinodina  milliaria,  R.  pachysperma,  and  Xanthoria  parie- 
titia  (figures  77,  78,  79,  80)  are  curious  in  that  the  four  have  rather 
different  distributions  outside  of  Long  Island.  It  would  seem  that  all  four 
have  a  high  degree  of  salt  tolerance  and  no  salt  requirement. 

3.  Vegetation  type.  An  understanding  of  the  distribution  of  lichens 
on  Long  Island  depends  on  a  knowledge  of  the  vegetation  types  in  which 
certain  lichens  are  most  likely  to  be  found.  That  is,  one  would  like  to 
know  which  lichens  are  “characteristic”  of  certain  vegetation  types.  In 
the  Braun-Blanquet  system,  a  characteristic  species  can  be  defined  by  its 
degree  of  “fidelity”  (see  Phillips,  1959).  Involved  in  the  analysis  of 
fidelity  is  an  intimate  knowledge  of  coverage,  sociability,  frequency,  and 
presence  derived  by  time-honored  methods  of  plot  studies  within  given 
stands. 

For  the  purposes  of  this  study  it  seems  best  to  use  collection  data 
as  a  source  of  information  concerning  the  distribution  of  lichens  in  vari¬ 
ous  vegetation  types,  considering  each  locality  to  be  a  “stand”  in  the 
Braun-Blanquet  usage.  In  these  collections,  all  vegetation  types  have 
been  represented  and  each  locality  essentially  has  a  complete  species  list. 
Since,  however,  within  a  given  locality  each  species  was  only  collected 
once  (p.  91)  and  without  any  notation  of  its  abundance,  cover,  soci¬ 
ability,  or  frequency,  only  “presence”  (the  percent  of  the  stands  of  a 
particular  vegetation  type  having  any  particular  lichen)  could  be  calcu¬ 
lated,  and  the  Braun-Blanquet  system  could  not  be  used  unaltered.  Know¬ 
ing  the  total  number  of  localities  in  which  each  species  was  collected  and 
the  number  of  collections  in  each  vegetation  type,  I  could  calculate  the 
percentage  of  its  occurrence  in  each  category,  calling  this  value  the 
vegetation  type  -  total  locality  value  or  VTL  value.  For  example,  Bacidia 
chlorococca  was  collected  in  58  different  localities  on  Long  Island. 
Four  of  these  were  dune  localities.  Since  there  are  22  dune  localities 
on  Long  Island,  the  presence  value  of  B.  chlorococca  in  dune  locali¬ 
ties  is  4/22  or  23  percent.  Since  there  are  58  specimens  of  B.  chlo¬ 
rococca  representing  58  localities,  the  VTL  value  of  the  species  in  dune 
localities  is  4/58  or  7  percent. 

If  a  species  has  a  relatively  high  VTL  value  in  the  same  vege¬ 
tation  type  for  which  it  has  a  high  presence  value,  it  can  be  called  “faith¬ 
ful"  in  much  the  same  context  as  fidelity  is  used  in  the  Braun-B'anquet 
system.  In  practice,  faithful  species  were  selected  by  a  nonmathematical 


270  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

system  of  examining  the  VTL  and  presence  values  (table  9)  and  select¬ 
ing  those  species  which  showed  particularly  high  values  in  only  one  (or 
sometimes  two  adjacent)  vegetation  types.  ("High  values”  were  con¬ 
sidered  to  be  a  minimum  of  20  percent  presence  plus  20  percent  VTL 
value.)  As  in  the  Braun-Blanquet  system,  faithful  species  often  are  rela¬ 
tively  uncommon  (Phillips,  1959). 

In  table  9,  presence  and  VTL  values  were  calculated  for  all  species 
that  occurred  in  10  or  more  of  the  135  stands  studied  in  Suffolk  County. 
Nassau  County  localities  were  excluded  because  of  the  possibility  of 
disturbance  due  to  the  proximity  of  New  York  City.  All  species  known 
to  be  bog-inhabiting  were  considered,  whether  or  not  they  occurred  in  ten 
stands,  since  there  were  only  eight  bogs  studied. 

In  order  to  make  the  numbers  of  the  various  vegetation  types  more 
directly  comparable,  some  categories  were  considered  together.  It  seemed 
valuable,  however,  to  separate  the  white  cedar  swamp  localities  from  the 
maple  swamps.  Roadside  trees,  cherry-locust  stands,  beech  climax  forests, 
all  of  which  were  very  infrequent,  and  a  few  localities  which  were  in¬ 
adequately  described  for  classification  were  excluded. 

Table  9  clearly  shows  that  most  species  do  not  have  a  narrow  re¬ 
striction  to  a  single  vegetation  type,  but  rather  are  distributed  more 
broadly  along  a  continuum  with  a  moderate  peak  in  one  (or  sometimes 
two)  categories.  A  number  of  species  bridge  the  gap  evenly  between  two 
categories  (e.g.,  Cladonia  clavulifera  between  downs  and  pine-oak  forests, 
Cladonia  capitata,  Parmeliopsis  placorodia,  Pertusaria  trachythallina,  and 
Physcia  stellaris  between  pine-oak  and  scarlet-black  oak  forests,  and 
Pertusaria  tuberculifera  between  scarlet-black  and  red  oak  forests.  In  a 
previous  paper  ( Brodo,  1961a),  I  discussed  in  detail  the  distribution  of 
some  common  Long  Island  lichens  along  a  pine-oak  to  red  oak  forest 
continuum. 

Within  each  vegetation  type  the  following  species  may  be  thought  of 
as  more  or  less  “characteristic”  as  defined,  from  the  Braun-Blanquet 
school,  by  Phillips  (1959).  Each  species  listed  below  is  given  a  fidelity 
rating  based  on  Phillips’  definitions  of  the  Braun-Blanquet  fidelity  cate¬ 
gories.  Category  5  is  made  up  of  species  occurring  almost  exclusively  in 
one  vegetation  type;  category  4  comprises  species  having  both  presence 
and  VTL  values  in  one  vegetation  type  more  than  twice  those  in  any 
other;  category  3  comprises  all  other  faithful  species  ( p.  269). 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  271 


Dunes,  downs,  and  sand  plains:  Pine  barrens,  pine-oak  forest: 


Acarospora  fuscata 

(4) 

Bacidia  chlorococca 

(3) 

Cetraria  islandica 

Cladonia  atlantica 

(4) 

subsp.  crispa 

(3) 

C.  clavulifera 

(3) 

Cladonia  boryi 

(4) 

C.  subtenuis 

(3) 

C.  clavulifera 

(3) 

Lecidea  anthrocophila 

(3) 

C.  furcata 

(4) 

L.  sc  alar  is 

(3) 

C.  strepsilis 

(4) 

L.  uliginosa 

(4) 

:arlet-black  oak  forest: 

Parmelia  galbina 

(3) 

Buellia  stillingiana 

(3) 

Parmeliopsis  placorodia 

(3) 

Lecanora  caesiorubella 

Pertusaria  trachythallina 

(3) 

subsp.  lathamii 

(3) 

Physcia  aipolia 

(4? 

Parmelia  subaurifera 

(3) 

Ph.  stellaris 

(3) 

Parmeliopsis  placorodia 

(3) 

Red  oak  forest: 

Pertusari a  trachythallina 

(3) 

Cladonia  coniocraea 

(3) 

P.  tuberculifera 

(3) 

Graphis  scripta 

(3) 

Pkyscia  stellaris 

(3) 

Lecidea  albocaerulescens 

(5) 

Pyrenula  nitida 

(4) 

Pertusaria  tuberculifera 

(3) 

rhite  cedar  swamp: 

Maple  swamp: 

Alectoria  nidulifera 

(3) 

(no  faithful  species) 

Cetraria  ci Haris 

(4) 

C.  viridis 

(5) 

Usnea  trichodea 

(4) 

Since  the  different  vegetation  types  occupy  different  portions  of  the 
island,  distribution  patterns  of  characteristic  species  and  others  close  to 
this  designation  reflect  their  specificities  in  maps  of  their  distribution  on 
Long  Island.  Distribution  maps  of  some  of  these  species  are  presented  in 
figures  33  to  63  and  74  to  81. 

a.  Dunes,  downs ,  and  sand  plains.  Distributions  of  lichens  strongly 
associated  with  dunes  and  sand  plains  are  mapped  in  figures  74  to  76. 
Cladonia  submitis  and  C.  uncialis  also  occur  in  open  sandy  or  grassy 
areas  within  the  scarlet-black  oak  forest  localities. 

b.  Pine  barrens  and  pine-oak  forests.  Those  species  more  or  less 
restricted  to  the  pine  forests  of  central  Long  Island  appear  to  be  either 
pine  specific  (figures  42  to  45)  or  confined  to  well-illuminated  oaks  (fig¬ 
ures  46  to  47  and  p.  267).  The  two  terricolous  lichens  characteristic  of 
this  vegetation  type  (Cladonia  calycantha  and  C.  floridana,  figures  48, 
49)  are  rather  narrowly  confined  to  very  acid  sand  in  open,  well-lighted 
localities  such  as  would  be  found  in  pine  forests.  Lecidea  varians,  Par- 
melia  galbina,  and  P.  perforata  (figures  50  to  52), although  basically  pine- 
oak  forest  species,  extend  eastward  in  well-illuminated  mixed  oak  and 
pine  stands  within  the  mature  oak  forest  region. 

c.  Morainal  (scarlet-black  oak  and  red  oak  forests).  Two  vegetation 
types  lie  along  the  glacial  moraines:  the  red  oak  forest  and  the  scarlet- 
black  oak  forest.  Gravelly,  sandy  loam  and  the  presence  of  many  boulders 
and  stones  characterize  both  vegetation  types,  and  consequently  many 
terricolous  and  saxicolous  species  are  distributed  along  one  or  both  of 
the  moraines.  Figures  53  to  56  include  the  terricolous  species,  figures  57 


272  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


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LICHENS  OF  LONG  ISLAND,  NEW  YORK 


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VEGETATION  TYPE  —  TOTAL  LOCALITY  ( VTL)  VALUE  FIDELITY 


274  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


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LICHENS  OF  LONG  ISLAND,  NEW  YORK 


275 


to  60  include  saxicolous  species,  and  figures  61  to  63  comprise  corticolous 
species.  For  more  detailed  ecological  notes,  the  reader  should  consult 
the  species  discussions  in  the  annotated  list. 

d.  Bogs  and  swamps.  The  cool,  humid  climate  of  a  bog,  together 
with  its  very  acid  soil  (or  water)  make  it  a  unique  habitat  with  a  rather 
unique  lichen  vegetation.  Maps  of  the  typical  bog  and  swamp  distribu¬ 
tions  should  be  compared  with  the  distribution  map  of  the  bog  and 
swamp  localities  (figure  32). 

Several  of  the  corticolous  bog  species  (figures  33,  34,  39)  are  con¬ 
fined  to  phorophytes  which  themselves  are  confined  to  bogs.  In  these  cases 
it  is  difficult  to  separate  substrate  influence  from  climatic  influence  in 
determining  the  causes  of  bog  specificity  (p.  267).  Climatic  effects  prob¬ 
ably  play  a  large  role  in  the  distribution  of  the  lignicolous  bog  lichens 
(figures  35  to  37)  as  well  as  those  bog-limited  species  also  found  grow¬ 
ing  on  trees  outside  the  bogs  (such  as  Pertusaria  amara  on  Acer  rubrum, 
figure  38,  and  Alectoria  nidulifera  on  Pinus  rigida.  figure  40). 

Parmelia  hypotropa  (figure  41)  shows  a  pattern  which  combines  bog 
and  swamp  localities  with  humid  maritime  localities,  a  distribution 
shared  by  Parmelia  perforata  (figure  52)  and  Ramalina  fastigiata. 

e.  Maritime.  Maritime  species  are  restricted  to  within  a  mile  of  the 
shore  (figures  77  to  81 )  and  are  within  the  aerohaline,  the  hygrohaline,  or 
the  hydrohaline  strata  of  the  shore  (p.  58).  Their  distributions  are 
probably  influenced  by  salt  water  or  spray,  high  winds,  high  illumination, 
or  some  combination  of  these  factors. 

FLORISTIC  CHANGES 

It  is  obvious  to  all  students  of  Long  Island  natural  history  that  the 
flora  and  fauna  of  the  island  have  significantly  changed  during  the  past 
50  to  75  years  and  are  still  changing.  One  need  only  list  the  lichens  col¬ 
lected  in  the  Brooklyn-Queens  area  prior  to  1900  to  see  a  striking 
example  of  these  changes.  Most  of  the  following  species  were  collected 
by  G.  B.  Brainerd  and  George  Hulst  in  1860’s. 


A  lectoria  nidulifera 
Anaptychia  palmulata 
Candelaria  concolor 
Cetraria  ciliaris 
C.  tuckermanii 
Cladonia  alpestris 
C.  bacillaris 
C.  capitata 
C.  chlorophaea 
C.  conista 
C.  cristatella 
C.  farinacea 
C.  f areata 
C.  pyxidata 


Cladonia  scabriuscula 
C.  submitis 
C.  sub  tenuis 
C.  uncialis 
C.  verticillata 
Collema  subfurvum 
Graphis  scripta 
Haematomma  sp. 
Lecanora  conizaea 
Leptogium  cyanescens 
Lobaria  pulmonaria 
L.  quercizans 
Pannaria  lurida 
Parmelia  aurulenta 


Parmelia  caperata 
P.  galbina 
P.  perforata 
P.  reticulata 
P.  stenophylla  (?) 

P.  sulcata 
Pcltigera  aphthosa 
P.  polydactyla 
Pertusaria  tuberculifera 
Physcia  millegrana 
Usnea  strigosa 
Xanthoria  fallax 
X.  parietina 


276 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Eastern  Long  Island  has  also  seen  some  alterations  in  the  flora,  but 
here  the  cause  has  mainly  been  the  great  hurricanes  of  1938  and  1944. 
Roy  Latham  (in  letter.  May  29,  1960),  describing  the  effects  of  these 
hurricanes  on  Long  Beach  at  Orient  Point  wrote  “Salt  water  flooded  all 
of  this  beach  which  was  exposed  to  gales  and  rolling  waves  and  the  beach 
was  swept  as  clean  as  a  new  house  floor.  In  places  the  water  was  four  to 
six  feet  in  depth  and  washed  the  bark  lichens  from  the  low  cedar  trunks 
and  wrenched  the  branch-growing  species  away.  All  traces  of  Usneas  and 
Ramalinas  disappeared  in  the  storm.  I  don’t  think  these  two  species 
have  appeared  there  since.  The  Cladonias  showed  a  fair  comeback  in 
two  years,  but  not  in  the  abundance  or  large  growth  of  the  old  days. 
I  know  that  alpestris  has  not  returned,  and  I  don’t  believe  rangiferina, 

| or]  sylvalica  [arbuscula]  may  have  returned.  After  the  second  hurricane 
of  1944,  the  beach  was  again  washed  by  high  flood  tides  and  left  about 
the  same  condition  as  in  1938.”  Latham  also  mentioned  local  building 
projects  and  farm  clearance  as  having  removed  the  last  stations  of  a  few 
of  the  rarer  species. 

It  is  not  only  this  generation,  however,  which  has  seen  the  gradual 
disappearance  of  lichens.  Willey  (1892,  p.  3),  reflecting  on  his  30  years 
of  collecting  in  New  Bedford,  Massachusetts,  wrote  "Of  late  years,  the 
clearing  of  forests,  the  quarrying  of  ledges,  and  the  breaking  up  of 
boulders,  have  tended  to  the  destruction  of  Lichens.  The  largest  of  the 
cypresses  (Chamaecy paris  thyoides)  have  gone  to  the  migratory  steam 
sawmill;  the  beeches  went  to  the  plane  factory;  and  the  hollies,  once 
abundant,  were  converted  into  knick-knacks,  so  that  few  of  any  size 
remain;  while  the  rocks  and  boulders  exist  only  in  the  foundations  of 
houses  and  factories. 

Of  the  total  lichen  flora,  47  species,  most  representing  Latham 
material,  have  not  been  collected  in  the  course  of  my  own  field  work.  Of 
these,  24  are  represented  by  only  one  specimen.  Several  of  the  remaining 
23  species,  perhaps,  are  actually  becoming  extinct  on  the  island.  Some 
mislabelling  is  involved  in  the  Latham  collection,  and  a  few  of  the  “rare” 
specimens  from  Latham’s  herbarium  may  actually  represent  material  sent 
to  Latham  on  exchange  (especially  from  northwestern  United  States) 
which  became  misplaced  and  then  mislabelled. 

Some  of  the  species  which  I  did  not  find,  together  with  some  col¬ 
lected  more  frequently  in  the  past  than  during  this  study  are  listed  below, 
followed  in  each  case  respectively  by  the  number  of  old  and  recent 
collections. 


Caloplaca  pyracea  (5-0) 
Cladonia  beaumontii  (21-1) 
C.  cdpestris  ( 1 4-2 ) 

C .  rangiferina  (25-2) 
Collema  subfurvum  (6-0 ) 
Lobaria  puhnonaria  ( 13-4) 
Pcltigera  praetextata  (19-5) 


Ramalina  stenospora  (7-0) 

R.  willeyi  (9-3) 

Teloschistes  chrysophthalmus  (9-0) 
T.  ftavicans  (2-0) 

Umbilicaria  muhlenbergii  (3-0) 
Verrucaria  silicicola  (9-2) 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


277 


Special  efforts  were  made  to  locate  all  these  species,  particularly  the 
two  Teloschistes,  the  Umbilicaria,  Cladonia  rangiferitia,  and  C.  alpestris. 
It  is  interesting  to  observe  that  for  most  of  these  species  Long  Island 
represents  the  outer  edge  of  their  natural  distribution.  It  is  the  southern 
limit  or  near-limit,  compared  with  areas  of  similar  altitude,  for  Cladonia 
alpestris,  Caloplaca  pyracea,  and  Umbilicaria  muhlenbergii;  it  is  the 
northern,  or  close  to  the  northern  limit,  for  Ramalina  stenospora,  Telo¬ 
schistes  chrysophthalmus,  and  T.  flavicans.  The  drainage  and  filling  of 
local  bogs  and  cutting  of  humid  forests  (particularly  affecting  the  popu¬ 
lations  of  oceanic  species  such  as  Lobaria  pulmonaria  and  Collema  sitb- 
furvum),  hurricanes,  building  projects,  and  environmental  pollution 
undoubtedly  all  took  their  toll.  Once  an  outlying  population  is  cut  down 
by  any  of  these  factors,  its  chances  of  re-expansion  are  much  slimmer 
than  those  of  species  lying  well  within  their  normal  or  potential  range. 
The  main  reasons  seem  to  be  that  these  marginal  populations  are  not 
living  under  optimum  conditions  for  reproduction,  and  the  chances  of 
reinvasion  are  small  due  to  low  population  levels  in  neighboring  areas. 

There  were  70  species  that  were  collected  only  in  my  own  field 
work.  The  great  majority  of  these  are  crustose  species  and  were  probably 
overlooked  by  previous  collectors.  It  would  be  foolish  to  attempt  a  general 
analysis  to  find  which,  if  any,  of  the  70  species  might  have  actually 
become  established  on  the  island  within  the  last  50  years.  It  is  possible 
to  state,  however,  that  Lecanora  muralis  is  apparently  an  adventive  on 
Long  Island,  probably  having  been  brought  in  on  limestone  building 
materials  from  some  area  to  the  north  or  west  where  both  limestone  and 
the  species  are  abundant. 


Summary  and  Conclusions 

HABITAT  ECOLOGY  AND  LICHEN  COMMUNITIES 

1.  Substrate  factors,  including  texture,  moisture-holding  capacity, 
stability,  and  chemical  composition  are  important  in  defining  lichen 
distributions  and  community  composition. 

2.  The  relationship  between  climatic  factors  (especially  rate  of 
fluctuation  and  degree  of  atmospheric  humidity)  and  available  light 
are  also  important. 

3.  If  some  of  the  characteristics  of  a  substrate  are  altered  by 
natural  or  unnatural  means,  the  lichen  communities  inhabiting  the  sub¬ 
strate  also  will  become  altered.  In  this  way,  two  substrates  of  basically 
unrelated  origin  may  bear  similar  floras  if  the  substrate  characteristics 
converge  and  become  close  to  identical. 

4.  Certain  aspects  of  substrate  characterizations,  especially  bark 
moisture  capacity,  were  discussed  in  detail.  No  method  for  expressing 
moisture  capacity  now  in  usage  appears  to  be  entirely  satisfactory. 

5.  The  actual  factors  involved  in  producing  communities  commonly 
called  “neutrophilic,”  “nitrophilic,”  or  “coniophilic”  need  much  more 
investigation  and  clarification  before  the  terms  can  be  used  in  a  mean¬ 
ingful  way.  In  most  cases,  it  is  similarly  difficult  to  distinguish  between 
“skiophilous”  and  “hygrophilous”  tendencies  in  lichens.  The  terms  should 
be  considered  as  mere'y  suggestive,  and  not  necessarily  reflecting  absolute 
eco'ogical  requirements. 

6.  The  results  of  transplant  experiments  used  in  the  study  of  ver¬ 
tical  lichen  distribution  on  tree  trunks  suggest  that  with  some  species 
(e.g.,  Cladonia  chlorophaea)  microclimate  is  the  limiting  factor,  and  in 
others  (e.g.,  Leccinora  caesiorubella) ,  it  is  some  aspect  of  the  substrate, 
or  competition,  that  is  limiting.  The  technique  of  transp'anting  bark  disks 
bearing  lichen  thalli  promises  to  be  important  in  studies  of  lichen  ecology. 

7.  The  lichens  of  red  oak  forests  were  sampled  in  an  east-west 
transect,  and  the  data  subjected  to  statistical  analysis.  From  these  results, 
certain  observations  were  made  concerning  species  composition  in  oak 
forests.  These  were  compared  with  the  results  of  an  earlier  sampling  of 
oak  and  pine-oak  forests  in  central  Long  Island,  and  conclusions  were 
drawn  pertaining  to  the  differences  in  lichen  flora  seen  in  the  two  vege¬ 
tation  types. 

8.  Continua  were  described  with  communities  on  tree  trunks  (fol¬ 
lowing  bark  age  and/or  vertical  position  gradients),  and  with  terricolous 
communities  (following  soil  type  gradients). 

9.  Lichen  successions  involving  corticolous,  terricolous,  and  saxico- 
lous  species  were  observed  and  described.  A  detailed  description  of  a 
primary  old  field  succession  was  presented.  Nondirectional  or  cyclic 
changes  involving  corticolous  and  dune-inhabiting  communities  were 
also  described. 


279 


280  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

10.  Lichen  communities  were  considered  to  be  groups  of  species 
living  under  similar  conditions  due  to  similarities  in  their  habitat  require¬ 
ments  and  tolerances  and  with  relatively  little  species  interaction. 

LICHEN  DISTRIBUTIONS 

1.  The  distribution  of  some  lichens  on  Long  Island  is  heavily  influ¬ 
enced  by  their  substrate  specificity,  and  of  others,  by  climatic  require¬ 
ments. 

2.  Most  species  on  Long  Island  are  more  or  less  confined  to  a 
particular  segment  of  the  forest  continuum,  usually  including  more  than 
one  vegetation  type. 

3.  Certain  vegetation  types  have  “characteristic”  lichen  species, 
just  as  they  have  characteristic  flowering  plants.  These  characteristic 
lichens  were  listed  and  discussed. 

4.  Based  on  a  sample  of  81  percent  of  the  total  lichen  flora,  21  per¬ 
cent  of  the  species  have  an  Arctic-boreal  distribution,  71  percent  are 
Temperate,  and  8  percent  are  Tropical.  In  addition,  24  percent  of  the 
species  are  endemic  to  North  America,  53  percent  are  circumboreal, 
11  percent  are  found  in  Europe  and  not  in  Asia  (almost  all  of  which 
are  amphiatlantic) ,  and  7  percent  are  found  in  Asia  and  not  in  Europe 
(almost  all  of  which  have  the  classic  Eastern  Asia-Eastern  United  States 
disjunct  pattern). 

5.  Origins  of  the  various  distributional  types  were  suggested,  and 
possible  migration  routes  to  Long  Island  were  outlined. 

6.  The  “fog  belt”  in  the  Montauk  region,  together  with  Chamaecy- 
paris  bogs,  have  a  large  number  of  oceanic  and  boreal  species.  Most  of 
the  coastal  plain  species  are  closely  restricted  to  bogs,  sand  dunes  and 
sand  plains,  and  salt  spray  habitats,  and  many  follow  the  limits  of  the 
pine-oak  forest.  East  Temperate  and  North  Temperate  species  are  com¬ 
monly  centered  in  the  red  oak  forests  along  the  north  shore. 

THE  CITY  EFFECT 

1.  A  detailed  study  of  the  influence  of  the  New  York  City  atmos¬ 
phere  on  Long  Island  lichen  distribution  was  carried  out  using  collection 
methods,  statistical  analyses  of  forest  samples,  and  transplant  experiments. 

2.  A  city  influences  lichen  distribution  both  through  its  induced 
drought  conditions  and  its  production  of  toxic  atmospheric  materials, 
presumably  SOL.  for  the  most  part.  Pollution  affects  the  lichens  at  greater 
distances  from  the  town  centers  than  does  city-induced  drought.  These 
conclusions  were  stated  as  tentative,  pending  direct  measurements  of  both 
humidity  and  pollution  levels  in  conjunction  with  lichen  growth. 

3.  It  was  suggested  that  on  Long  Island  the  pollution  effect  is  so 
strong  that  most  corticolous  lichens  are  killed  at  a  distance  beyond 
which  the  drought  effect  can  influence  their  vertical  distribution. 

THE  LICHEN  FLORA 

1 .  Despite  its  comparatively  small  size.  Long  Island  shows  a  sur¬ 
prising  diversity  in  its  vegetation,  both  phanerogamic  and  cryptogamic. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


281 


2.  Including  only  material  personally  investigated,  261  species 
in  65  genera  and  28  families  were  cataloged.  Literature  records  were 
excluded  from  the  list  because  of  the  high  frequency  of  misidentifications 
and  recent  reinterpretations  of  many  species. 

3.  Three  names  were  introduced  as  new  to  science:  Polyblastiopsis 
quercicola,  Pertusaria  subpertusa,  and  Lepraria  zonata.  The  Lepraria 
had  a  previous  but  invalid  description. 

4.  The  following  are  documented  for  the  first  time  from  North 
America,  and  have  been  listed  in  the  latest  North  American  checklist 
(Hale  and  Culberson,  1966): 

Arthonia  mediella  Pertusaria  tuberculifera 

A.  sexlocularis  Porina  hibernica 

Ochrolechia  parella 

5.  The  following  names  from  the  Long  Island  lichen  flora  are  not 
included  in  the  North  American  checklist  (ibid): 

Bacidia  “intermedia”  Pseudevernia  furfuracea 

lonaspis  odora  Stereocaulon  saxatile 

Lecanora  degelii 
Lecidea  aeruginosa 


Appendix  A 

LONG  ISLAND  COLLECTORS 


Name 

Alexander,  E.  J. 

Ames,  F.  H. 

Austin,  Maud  G. 
Booth,  M.  A. 
Brainerd,  George  B. 

Britton,  E.  G. 
Brodo,  Irwin  M. 
Cain,  Stanley  A. 

Carnegie,  T.  M. 
Clute,  Willard  N. 
Copeland,  Joseph  J. 
Culberson,  W.  L. 
Davis,  William 

Dillman,  George 

Gillis,  W.  T. 

Grier,  N.  M. 

Grout,  A.  J. 

Harper,  R.  M. 

Harris,  A.  E.  G. 
Hulst,  George  D. 
Imshaug,  H.  A. 
Latham,  Roy 


Lloyd,  F.  E. 
Morgan,  D.  P.  J.  M. 


A  ppro.x. 

dates  Herbarium 


1926 

May  1910 

NYS 

? 

BKL 

1877 

FH 

1860-1866 

BKL 

1897-1898 

NY 

1 959-present 

NYS.MSC 

1 930’s 

NY 

1913-1914 

FH : Howe 

1898 

NY 

1 940’s 

MSC 

1 950’s 

FH 

1912-1929 

Staten 

Island 

1927 

NY: 

Torrey 

1961 

MSC 

1  900 

BKL 

1918 

NY 

1904 

MICH 

1890 

BKL 

1960 

MSC 

1  908-present 

CUP,N  YS 
FH,MO, 
MICH, 
LATHAM 

MSC 

1896 

NY 

1909 

FH: Howe 

A pprox. 
localities 
Bellmore, 

High  Hill  Beach 
Brookhaven 
“Long  Island” 

Orient 

New  York  City 
and  vicinity 
Sag  Harbor 

Throughout  Long  Island 
Selden, 

“Cold  Spring  Harbor” 
Massapequa 
Southampton 
Southampton 
Montauk  region 
Riverhead  region 

Yaphank,  Farmingville, 
Wading  River 

Orient  Pt. 

Montauk 

“Cold  Spring  Harbor” 
Cold  Spring  Harbor 
and  vicinity 
Meadowbrook  Valley; 
Hempstead  PI. 

Cold  Spring 
New  York  City 
Eastern  Long  Island 


Eastern  Long  Island 

Sayville 

Southampton 


282 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


283 


Name 

Approx. 

dates 

Herbarium 

A  pprox. 
localities 

Ogden,  Eugene  C. 

1 950-present 

NYS 

Eastern  Long  Island 

Peck,  Charles  H. 

1 860's-l  914 

NYS 

Throughout  Long  Island 

Schrenk,  Hermann 

1894 

MO 

Eastport 

Schrenk, Joseph 

? 

NY 

College  Pt.,  Jamaica 

Smith,  Stanley  J. 

1 950’s-present 

NYS 

Eastern  Long  Island 

Taylor,  Norman 

1918-1922 

BKL 

East  Point,  Coram 

Torrey,  Raymond 

1 930's 

NY 

Throughout  Long  Island 

Von  Scheur 

1895 

MO 

Montauk  Point 

Warner,  E.  A. 

1 900 

BKL 

Valley  Stream 

Young,  Alfred  R. 

9 

BKL 

Orient 

Appendix  B 

GLOSSARY 

I.  MORPHOLOGICAL  AND  ECOLOGICAL  TERMS 

The  emphasis  in  this  portion  of  the  glossary  is  on  morphological 
terminology  as  used  in  lichenology  and  with  definitions  designed  to  aid 
experienced  observers  as  well  as  individuals  with  little  or  no  background 
in  mycology  or  lichenology.  No  attempt  was  made  to  include  all  eco¬ 
logical  terminology,  since  most  terms  were  defined  or  explained  in  the 
text  when  they  were  used.  Some  ecological  terms  of  special  importance 
in  the  identification  of  lichens  were  included,  however.  Chemical  termi¬ 
nology  is  treated  in  part  II  of  this  glossary. 

Acicular.  Needle-shaped,  i.e.,  slender  and  pointed  at  both  ends. 

Adnate.  Closely  attached  to  a  surface,  with  few  or  no  ascending  parts. 
Amphigymnioid.  In  foliose  lichens,  lacking  rhizines  close  to  the  edges 
of  the  lower  surface  although  having  rhizines  in  the  center,  as  in 
the  subgenus  Ainphigymnia  of  the  genus  Parmelia. 

Amphithecium.  The  portion  of  a  lecanorine  apothecium  external  to  the 
proper  exciple  (figure  87A),  usually  containing  algae;  the  thalline 
margin. 

Ampliariate.  In  Pertusaria ,  pertaining  to  fruit  warts  which  are  broadest 
at  the  base,  as  in  the  subgenus  Ampliaria. 

Anisotomic  branching.  In  Cladonia,  especially  the  subgenus  Cladina, 
unequal  branching  which  results  in  a  more  or  less  distinct  main  axis 
from  which  smaller,  more  slender  branches  arise. 

Apical.  At  the  apex  or  tip  of  a  stalk  or  lobe. 

Apothecium.  A  disk-  or  cup-shaped  ascocarp  (figure  87). 

Areolate.  Broken  up  into  small,  irregular,  usually  angular  patches  (are- 
oles),  often  appearing  tile-like. 

Articulated.  Divided  into  short  or  long  segments  and  having  conspicuous 
joints. 

Ascocarp.  The  fruiting  body  of  an  Ascomycete;  the  structure  which  bears 
the  asci  which  in  turn  contain  the  ascospores. 

Ascohymenial.  Pertaining  to  a  type  of  ascocarp  having  true  paraphyses 
and  unitunicate  asci;  characteristic  of  the  subclass  Ascomycetidae. 
Ascolocular.  Pertaining  to  a  type  of  ascocarp  in  which  the  asci  (gen¬ 
erally  bitunicate)  arise  within  a  uniform  stromatic  mass  and  are 
separated  in  maturity,  not  by  true  paraphyses,  but  by  paraphysoid 
threads;  characteristic  of  members  of  the  subclass  Loculoasco- 
mycetidae. 

Ascospore.  A  spore  produced  in  an  ascus. 

Ascus  (asci).  The  sac-like  structure  in  Ascomycetes  in  which  the  asco¬ 
spores  are  formed. 

Aspicilioid.  Having  apothecia  sunken  into  the  thallus  so  that  the  apo- 

284 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  285 

thecial  disk  is  level  with  the  thallus  surface  or  slightly  depressed;  as 
in  the  section  Aspicilia  of  the  genus  Lecanora. 

Axil.  In  Cladonia  thalli,  the  point  at  which  two  or  more  branches  or  a 
branch  and  the  main  axis  meet. 

Axis,  (a)  The  main  trunk  or  stem  of  an  abundantly  branching  thallus. 
(b)  In  Usnea,  the  cartilaginous  (chondroid)  central  core  running 
through  the  thallus  filaments. 

Bacilliforin.  Rod-shaped  and  generally  very  small. 

Biseriate.  Spores  in  two  rows  within  the  ascus. 

Branching  ( di tri-,  tetrachotomy) .  In  Cladonia.  especially  Cladina.  refers 
to  the  number  of  equal  branches  coming  off  at  any  one  axil  (two, 
three,  and  four  respectively). 

Caespitose.  Tufted;  shrubby. 

Calcareous  rock.  Rock  containing  lime  and  producing  vigorous  bubbling 
(CCL,)  upon  application  of  a  strong  acid. 

Canals.  In  some  Pertusaria  spores,  fine  lines  or  channels  on  or  within  the 
outer  or  inner  spore  walls,  and  communicating  with  the  spore  lumen. 

Capitate.  Having  a  rounded  or  “head-like”  shape,  usually  referring  to  a 
type  of  soralium. 

Carbonaceous.  Opaque  black,  and  usually  brittle. 

Cartilaginous.  Referring  to  tissues  which  are  transluscent  and  somewhat 
stiff;  chondroid. 

Cephalodium  (cephalodia) .  A  small  gall-like  growth  occurring  in  large 
numbers  within  the  tissues  or  on  the  surfaces  of  some  lichens;  gen¬ 
erally  containing  blue-green  algae. 

Channelled.  Referring  to  spore  wall  markings  in  Pertusaria  (see  canals). 

Clunky.  Minutely  and  irregularly  cracked. 

Chondroid.  See  Cartilaginous. 

Cilia.  Hair-like  thalline  appendages;  occurring  at  the  thallus  or  apothecial 
margins  of  many  foliose  and  fruticose  lichens. 

Cinereous.  Grey-ashy  in  color. 

Clavate.  Club-shaped;  i.e.,  broader  at  one  end  than  the  other. 

Continuous.  Thallus  unbroken,  or  broken  very  little,  by  cracks. 

Coralloid.  (a)  Having  or  being  composed  of  small,  minutely  branched 
cylindrical  outgrowths,  (b)  A  type  of  isidium  having  this  form. 

Cortex.  The  outer  protective  layers  of  a  lichen  thallus  or  apothecium; 
completely  fungal  in  composition;  often  cellular  in  appearance 
(paraplectenchymatous),  but  may  have  other  forms  as  well  (figure 
87A) . 

Corticolous.  Growing  on  bark. 

Crenate.  Having  a  margin  with  rounded  teeth  or  minute  lobes. 

Crenulate.  Finely  crenate. 

Crustose.  A  thallus  type  which  is  generally  in  contact  with  the  substratum 
at  all  points  and  lacks  a  lower  cortex;  cannot  be  removed  intact  from 
its  substrate  without  removing  a  portion  of  the  substrate  as  well. 

Cyanophyceaen.  Pertaining  to  blue-green  algae. 


286  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Decorticate.  Having  had  a  cortex  which  has  now  fallen  away  or  decom¬ 
posed. 

Dicarpous.  With  two  ascocarps;  usually  refers  to  two  apothecia  per  fruit 
wart  in  species  of  Pertusaria. 

Dichotomy.  See  Branching. 

Disk.  The  flat,  concave,  or  convex  surface  of  an  apothecium;  usually 
pigmented  in  a  characteristic  way. 

Dispersed.  Pertaining  to  a  thallus  which  consists  of  scattered  small  areoles 
or  granules. 

Dorsi-ventral.  With  recognizable  upper  and  lower  surfaces. 

Ecorticate.  Never  having  had  a  cortex. 

Effigurate.  Referring  to  the  lobed  margin  of  a  thick,  basically  crustose 
thallus. 

Effuse.  Pertaining  to  a  thallus  having  no  distinguishable  boundaries. 

Ellipsoid.  Oval  to  elongate-oval  in  outline. 

Endolithic.  Growing  “within”  a  rock,  i.e.,  under  and  around  the  rock 
crystals,  often  with  little  or  no  thallus  visible  on  the  outer  rock 
surface. 

Epilithic.  Growing  on  a  rock  surface  with  little  or  no  penetration  be¬ 
tween  and  under  the  rock  particles. 

Epiphloedal.  In  corticolous  lichens,  a  thallus  having  little  or  no  pene¬ 
tration  below  the  outermost  bark  layer  (figure  88B). 

Epispore.  A  transparent  gelatinous  covering,  often  irregular  in  thickness, 
surrounding  the  ascospores  of  many  lichens;  often  called  a  “halo.” 

Epithecium.  The  uppermost  portion  of  the  hymenium  formed  by  the 
expanded  tips  of  paraphyses;  usually  pigmented  and  sometimes  in- 
spersed  with  tiny  granules. 

Eupertusariate.  In  Pertusaria.  pertaining  to  fruit  warts  which  are  more 
or  less  constricted  at  the  base,  as  in  the  subgenus  Eupertusaria  of 
Erichsen. 

Exciple.  (a)  An  area  in  an  apothecium  external  to  and  below  the  hypo- 
thecium,  forming  the  apothecial  margin  in  lecideine  apothecia 
(figure  87B)  and  internal  to  the  amphithecium  in  lecanorine  apothecia 
(figure  87A).  The  "proper  exciple”  of  Fink  (1935  ).  Note:  Fink  con¬ 
sidered  only  the  area  lateral  to  the  hymenium  as  the  excip'e,  with 
the  portion  below  the  hymenium  being  called  the  “hypothecium.” 
The  hypothecium  as  used  here  refers  only  to  the  subhymenial  tissue 
above  the  excip'e,  (the  “subhymenium”  of  Degelius  [1954]). 
(b)  The  inner  wall  of  a  perithecium,  generally  circular  in  cross- 
section;  can  be  hyaline,  pigmented,  or  carbonaceous  (figure  87C). 

Excipuloid  tissue.  Tissue  forming  the  walls  or  margins  of  ascolocular 
ascocarps  (especially  in  Micarea  and  Arthonia)  similar  in  appear¬ 
ance  and  position  to  the  true  exciple  of  lecideine  apothe~ia. 

Falcate.  Bending  in  one  direction;  sickle-shaped. 

Farinose  soredia.  Very  fine,  powdery  soredia. 

Flexuous.  Bending  in  alternate  directions,  i.e.,  "zig  zag.” 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


287 


Fuliose.  Pertaining  to  a  more  or  less  “leafy”  lichen  thallus,  distinctly 
dorsi-ventral,  and  varying  in  its  attachment  to  the  substrate  from 
almost  completely  adnate  to  umbilicate. 

Fruit  wart.  In  Pertusaria,  a  thalline  wart  (verruca)  which  contains  one 
or  more  apothecia. 

Fruticose.  Pertaining  to  a  lichen  thallus  which  is  podetioid,  pendent, 
or  shrubby. 

Fusiform.  Narrow,  tapering  toward  both  ends,  usually  with  pointed  ends; 
spindle-shaped. 

Glabrous,  (a)  Having  a  more  or  less  smooth,  shiny  surface,  (b)  With 
no  trace  of  tomentum. 

Globose.  Nearly  spherical. 

Granular,  (a)  Having  granules  or  granule-like  particles,  (b)  Pertaining 
to  soredia,  composed  of  particles  large  enough  to  be  easily  distin¬ 
guished  under  a  dissecting  microscope,  presenting  a  coarse  appear¬ 
ance,  not  powdery  as  in  farinose  soredia. 

Granule,  (a)  In  thalli,  a  spherical  or  nearly  spherical  corticate  particle, 
(b)  Pertaining  to  chemical  materials,  any  small  regular  or  irregular 
particle,  opaque  or  hyaline,  found  associated  with  various  lichen 
tissues. 

Gyrose.  Having  a  folded  or  ridged  surface;  referring  to  apothecia,  par¬ 
ticularly  in  Umbilicaria,  which  show  the  invasion  of  concentric  or 
radiating  rows  of  sterile  excipular  tissue  into  the  hymenium. 

Halophytic.  Growing  in  habitats  having  high  salt  concentrations. 

Hyaline.  Colorless. 

Hygrophilous.  Generally  associated  with  moisture  (usually  high  atmos¬ 
pheric  humidity). 

Hymenium.  The  fertile  layer  of  an  ascocarp,  consisting  of  asci  and  para- 
physes,  or  paraphysoid  threads)  (figure  87). 

Hypha  ( hyphae ).  A  fungal  filament. 

Hypophloedal.  In  corticolous  lichens,  in  which  most  or  all  of  the  thalline 
tissue  is  below  one  or  more  layers  of  cork  (figure  88A). 

Hypothecium.  The  tissue  just  be'ow  the  hymenium  but  above  the  exciple 
(figure  87);  often  difficult  to  distinguish  from  the  exciple,  of  which 
some  authors  consider  it  a  part  (see  exciple). 

Hypothallus.  A  special  differentiated  hyphal  tissue  on  the  lower  surface 
of  some  lichens,  e.g.,  Anzia. 

Hypotrachynoid.  Having  rhizines  growing  over  the  entire  lower  thallus 
surface,  as  in  the  subgenus  Hypotrachyna  of  the  genus  Parmelia. 

Hysterothecium.  An  elongate  to  linear  ascocarp  seen  in  some  members  of 
the  Loculoascomycetidae,  e.g.,  Opegrapha. 

Imbricate.  Pertaining  to  sca'es  or  squamu'es  which  overlap  in  a  shingle¬ 
like  fashion. 

Inflated.  Swollen  and  hollow. 

Involucrellum.  The  exposed  covering  or  cap  external  to  the  excipulum 


288  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

present  on  many  perithecia;  usually  black  and  carbonaceous,  but  in 
some  species,  may  be  colorless  or  even  contain  algae  (figure  87C). 

Involute.  With  margins  rolled  inward. 

Isidiitm  ( isidia ).  A  minute,  cylindrical,  or  coralloid  thalline  outgrowth 
which  is  corticate  and  contains  algae;  apparently  functions  as  a 
vegetative  reproductive  body. 

Isotomic  branching.  Branching  into  sub-branches  of  equal  size,  resulting 
in  a  thallus  having  no  distinguishable  main  axis. 

Isthmus  ( isthmi ) .  The  narrow  canal  between  the  two  locules  of  a  polari- 
locular  spore  (figure  90A). 

Labriform.  (a)  Lip-shaped,  (b)  Pertaining  to  soralia,  generally  formed 
by  an  involute  thallus  margin  or  a  bursting  hollow  thallus  lobe, 
sorediate  on  the  lower  or  inside  (i.e.,  exposed)  surface,  as  in 
Hypogymnia  physodes. 

Lacinia  (laciniae) .  A  long,  slender  thallus  lobe,  as  in  Pseudevernia  and 
Ramalina. 

Laciniate.  Having  elongated,  narrow  lobes. 

Lamellate.  In  thin  plates  or  sheets. 

Laminal.  On  the  fiat,  usually  upper  surface  of  a  thallus. 

Lax.  Loose;  not  compact. 

Lecanorine.  Pertaining  to  an  apothecium  having  a  distinct  amphithecium, 
usually  containing  algae,  as  in  the  genus  Lecanora  (figure  87A). 

Lecideine.  Pertaining  to  an  apothecium  in  which  there  is  no  distinguish¬ 
able  amphithecium  and,  therefore,  in  which  the  exciple  forms  the 
apothecial  margin  (i.e.,  the  proper  margin),  as  in  the  genus  Lecidea 
(figure  87B). 

Lenticular.  Shaped  like  a  double  convex  lens. 

Leprose.  Composed  almost  entirely  of  loosely  organized  granules  or 
soredia. 

Lignicolous.  Growing  on  bare  wood  (lignum),  as  on  a  decorticate  log 
or  a  wooden  fence. 

Lirella  ( lirellae ).  An  elongate  to  linear  apothecium,  often  branched,  as 
in  Graphis. 

Lumen  ( lumina ).  A  cell  cavity,  occupied  by  the  protoplast. 

Macula  (maculae).  A  very  small  white  spot  or  blotch  on  the  surface  of 
a  thallus,  not  associated  with  any  break  in  the  cortex,  but  simply 
representing  a  locally  decolorized  or  alga-less  area. 

Maculiform.  (a)  Like  a  spot,  (b)  Referring  to  a  type  of  small,  rounded, 
laminal  soralium. 

Maritime.  Having  some  association  with  the  ocean. 

Mazaedium.  A  mass  of  ascospores  and  paraphyses  formed  by  the  disin¬ 
tegration  of  the  asci  of  a  special  type  of  ascocarp,  as  in  Chaenotheca. 

Medulla.  The  internal  region  in  a  thallus  or  lecanorine  apothecium  which 
is  generally  composed  of  loosely  packed  hyphae  (figure  87A). 

M ischoblastiomorphic .  Pertaining  to  a  spore  with  two  funnel-shaped 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


289 


Figure  87.  Ascocarps.  (a)  Lecanorine  apothecium,  as  in  thecium;  cor,  cortex;  epi,  epithecium;  exc,  exciple;  hym,  hy- 

Lecanora  spp.;  (b)  iecideine  apothecium,  as  in  Lecidea  spp.;  menium;  hyp,  hypothecium;  inv,  involucrellum;  med,  medulla; 

(c)  perithecium,  as  in  Porina  hibernica;  (d)  pseudothecium,  par,  paraphysoid  threads;  sti,  stipe;  str,  stroma, 
as  in  Polyblastiopsis  quercicola.  alg,  algal  layer;  arnph,  amphi- 


290 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


locules  (the  two  locules  appearing  like  an  hourglass  in  section) 
(figure  90B). 

Molariform.  Shaped  like  a  short,  blunt  tooth. 

Moniliform  cells.  Globose  hyphal  cells  joined  together  in  a  bead-like 
chain. 

Monocarpous.  Containing  one  apothecium. 

Murale.  Muriform. 

Muriform.  Having  both  longitudinal  and  transverse  septa,  with  the  cells 
thus  appearing  like  a  brick  wall  (figure  90D). 

Naked,  (a)  Pertaining  to  a  thallus  without  rhizines  on  the  lower  surface, 
(b)  Epruinose. 

Neutrophytic.  Growing  on  a  substrate  having  a  pH  close  to  7  (i.e.,  sub¬ 
strates  which  are  neither  distinctly  acid  nor  basic). 

Nitrophilous.  Showing  a  close  association  with  substrates  rich  in  nitrogen 
compounds. 

Nitrophobous.  Showing  a  distinct  disassociation  with  substrates  rich  in 
nitrogen  compounds. 

N itrophytic.  Showing  a  tendency  towards  being  nitrophilous. 

Nostoc.  A  genus  of  blue-green  algae  found  in  many  lichens;  producing 
bead-like  chains  or  filaments  when  free  living,  hut,  when  lichenized, 
may  be  single-  or  few-celled  (figure  89C). 

Octos porous.  Having  eight  spores  per  ascus. 

Orbicular.  Circular  in  outline. 

Ostiole.  The  small,  round,  apical  pore  in  various  types  of  perithecia, 
pseudothecia,  and  even  fruit  warts  of  Pertusaria. 

Pachyspore.  An  ascospore  with  uniformly  thickened  walls  and  spherical 
lumina  (figure  90C). 

Papilla  (papillae).  A  small,  generally  conical,  thalline  outgrowth,  having 
an  unbroken  cortical  covering. 

Paraphysis  (paraphyses) .  A  sterile  hypha,  sometimes  branched,  associated 
with  asci  in  the  hymenium  of  a  member  of  the  Ascomycetidae. 

Paraphysoid  threads  (or  filaments) .  The  remains  of  stromatic  tissue 
found  between  the  asci  in  ascolocular  ascrocarps;  often  is  highly 
branched  and  anastomosing. 

Paraplectenchymatous.  Pertaining  to  fungal  tissue  which  appears  cellular 
in  section  due  to  short  cells  and  a  highly  branched,  irregularly 
oriented  hyphal  system. 

Pellucid.  Almost  transparent. 

Peltate.  Attached  at  the  center  of  the  lower  surface. 

Pendulose.  Pendent;  hanging  down,  with  little  or  no  horizontal  or  erect 
growth. 

Perithecium.  A  flask-shaped  ascocarp  characteristic  of  members  of  the 
Sphaeriales  (figure  87C);  may  he  sessile,  or,  more  commonly,  sunken 
partially  or  completely  into  the  thallus  tissue. 

Phorophyte.  The  tree  or  shrub  upon  which  a  corticolous  lichen  is  growing. 

Phycobiont.  The  algal  component  (symbiont)  in  a  lichen  thallus. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


291 


Figure  88.  Thallus  types,  (a)  Hypophloedal;  (b)  epiphloedal.  bk,  bark; 
nec,  necrotic  layer  of  thallus;  thal,  living  thallus  tissue. 


10  20  30  40  50  /U. 

I  I  .-l - I  ...I  I 


Figure  89.  Lichen  phycobionts  (camera  lucida  drawings),  (a)  Trente- 
pohlia  (from  Graphis  scripta)\  (b)  Trebouxia  (from  Cladonia  sp.); 
(c)  Nostoc,  (front  Leptogium  cyanescens). 


Figure  90.  Some  ascospore  types,  (a)  Polarilocular;  (b)  mischoblastio- 
ntorphic;  (c)  pachysporous;  (d)  muriform. 


292  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

Phyllocladium  (phyllocladia) .  A  minute,  often  lobed  or  scale-like  out¬ 
growth  of  the  pseudopodetia  of  some  members  of  the  genus  Stereo- 
caulon. 

Platysmoid.  A  tissue  which  consists  of  “densely  agglutinated  thick-walled 
hyphae  with  very  narrow  lumina  .  .  (Dahl,  1952,  p.  129),  as  in 
the  subgenus  Platysma  of  the  genus  Cetraria. 

Podetioid.  Having  the  general  appearance  of  a  podetium. 

Podetium.  A  stalk  formed  by  a  vertical  extension  of  apothecial  tissues 
(usually  the  hypothecium  and  stipe);  the  stalk  usually  becomes 
secondarily  invested  with  an  algal  layer  and  cortex  (as  in  Cladonia) 
and  can  be  either  short  and  unbranched,  or  quite  tall  and  highly 
branched. 

Polarilocular.  Pertaining  to  spores  having  two  lumina  separated  by  a 
relatively  thick  septum  through  which  a  narrow  canal  or  isthmus 
passes  (figure  90A),  characteristic  of  members  of  the  Teloschistaceae. 

Polycarpous.  Two  or  more  apothecia  per  fruit  wart  (in  Pertusaria) . 

Polysporous.  More  than  eight  spores  per  ascus. 

Primary  squamide.  The  scale-like  component  of  the  primary  thallus  of 
a  Cladonia  species. 

Primary  thallus.  The  thallus  of  a  Cladonia  species  exclusive  of  the  podetia; 
generally  composed  of  leafy  scales  or  squamules,  but  sometimes  (as 
in  the  subgenus  Cladina)  composed  of  a  granular  crust. 

Proper  margin.  See  exciple. 

Prothallus.  The  non-assimilative  lower  portion  of  a  lichen  thallus  seen 
around  the  outer  edge  of  many  crustose  species  as  a  white  or  pig¬ 
mented  margin,  and  often  visible  as  a  mat  between  the  areoles  or 
granules  of  other  crustose  species. 

Pritinose.  Having  a  frosted  appearance  (usually  white  or  grey). 

Pseudocyphella  (pseudocyphellae) .  A  tiny  white  dot  or  pore  seen  in  large 
numbers  on  the  upper  and  sometimes  the  lower  thallus  surfaces  of 
many  foliose  species;  caused  by  a  break  in  the  cortex  and  the  exten¬ 
sion  of  medullary  hyphae  to  the  surface. 

Pseudopodetium  {pseudopodetia) .  A  podetioid  stalk  formed  by  a  vertical 
extension  or  growth  of  thalline  tissues;  like  true  podetia,  they  can  be 
simple  (as  in  Pycnothelia)  or  highly  branched  (as  in  Stereocaulon) . 

Pseudothalline  margin.  A  margin  of  thalline  origin  external  to  the  amphi- 
thecium  in  lecanorine  apothecia,  and  external  to  exciple  in  lecideine 
apothecia. 

Pseudothecium  (pseudothecia) .  The  ascocarp  of  a  member  of  the  Locu- 
loascomycetidae  which  appears  superficially  like  a  perithecium 
figure  87D). 

Punctiform.  Dot-like  and  very  minute. 

Pustulate.  Having  large  and  small  blister-like  protuberances  over  the 
thallus  surface,  each  blister  on  the  upper  surface  having  a  corre¬ 
sponding  depression  or  pit  on  the  lower  surface. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  293 

Pycnidial  jelly.  A  gelatinous  substance  found  in  the  pycnidial  cavity  of 
some  species  of  Cladonia. 

Pycnidium  (pycnidia).  A  globular  or  flask-shaped  body,  usually  very 
small,  in  which  pycnoconidia  are  formed;  often  closely  resembling 
a  perithecium  in  external  appearance;  the  “spermagonium”  of  many 
authors. 

Pycnoconidinm  (pycnoconidia) .  A  small,  spore-like  body  formed  in  a 
pycnidium;  apparently  can  act  as  a  conidium  (an  asexual  spore)  in 
some  species  and  a  spermatium  (a  type  of  male  gamete)  in  others; 
it  is  what  has  been  called  a  microconidium. 

Reniform.  Kidney-shaped. 

Reticulate.  Having  a  net-like  appearance  due  to  cracks,  pigmentation, 
ridges,  etc. 

Revolute.  Pertaining  to  margins  which  are  rolled  backward  or  downward. 

Rhizine  ( rhizinae ).  A  purely  hyphal  extension  of  the  lower  cortex,  which 
generally  serves  to  attach  a  foliose  thallus  to  its  substrate;  of  various 
lengths,  thicknesses,  colors,  and  degrees  of  branching. 

Rimose.  Having  a  minutely  cracked  appearance. 

Rugose.  Having  a  wrinkled  surface. 

Rugulose.  Having  a  minutely  wrinkled  surface. 

Saxicolous.  Growing  on  rock,  stone,  pebbles,  concrete,  or  brick. 

Scrobiculate.  Having  a  pitted  appearance. 

Scurfy.  Having  a  fine  powdery  or  scaly  surface  (not  synonomous  with 
sorediate ) . 

Septum  (septa) .  A  crosswall  in  a  hypha  or  spore. 

Sessile.  Without  a  stalk  of  any  kind. 

Sigmoid.  Shaped  like  an  “S.” 

Siliceous  rocks.  Rock  composed  mainly  of  silicon  compounds,  producing 
no  bubbles  upon  application  of  a  strong  acid.  Quartz  and  granite  are 
examples. 

Simple.  Unbranched. 

Skiophilous.  Showing  a  strong  association  with  shaded  habitats. 

Soralium  (soralia) .  A  body  or  area  in  which  soredia  are  produced;  can 
be  in  many  forms. 

Sordid,  (a)  Dark,  (b)  Appearing  “dirty.” 

Soredium.  A  vegetative  reproductive  body  of  a  lichen  consisting  of  a 
few  algal  cells  entwined  and  surrounded  by  a  layer  of  fungal  hyphae; 
entirely  ecorticate;  generally  produced  in  localized  masses  called 
soralia,  or  covering  large  diffuse  areas  of  a  thallus. 

Spore.  A  single-  or  few-celled  reproductive  body  capable  of  giving  rise 
to  a  new  plant;  as  used  here,  refers  specifically  to  an  ascospore. 

Squamiform.  Scale-  or  squamule-shaped. 

Squamule.  A  small,  scale-like  lobe  or  areole,  generally  at  least  partially 
ascending. 

Stipe.  In  apothecia,  the  central  stalk-like  extension  of  the  exciple  down¬ 
ward  and  into  the  thallus. 


294 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Stipitate.  Raised  on  a  stalk  or  stipe. 

Stramineous.  Straw-colored. 

Stroma.  A  closely-packed  mass  of  hyphae,  often  carbonaceous,  which  is 
generally  associated  with  reproductive  structures. 

Striate.  Having  a  longitudinally  striped,  grooved,  or  ridged  appearance. 

Strigose.  Bearing  dense,  short,  hair-like  projections  or  branches. 

Sub-  (a)  Partially,  (b)  Incompletely,  (c)  Approaching,  (d)  Under. 

Subcanaliculate.  With  shallow  channels  or  furrows. 

Subfoliose.  Pertaining  to  a  crustose  species  with  marginal  lobes,  showing 
some  tendency  towards  becoming  ascending. 

Substrate.  The  material  upon  which  a  lichen  is  growing  or  to  which  it 
is  attached. 

Subulate.  Elongate,  and  gradually  tapering  to  a  point. 

Terete.  Circular  in  cross  section. 

Terricolous.  Growing  on  soil  or  sand. 

Tetrachotomy.  See  branching. 

Thalline.  Pertaining  to  the  lichen  thallus;  similar  to  the  thallus  in  appear¬ 
ance  or  structure. 

Thalline  margin.  See  amphithecium. 

Thallus.  In  lichens,  the  vegetative  plant  body  consisting  of  both  algal 
and  fungal  components. 

Tier.  A  platform-like  expansion  on  the  podetia  of  several  species  of 
Cladonia  (e.g.,  Cladonia  verticillata)  at  which  point  one  or  more 
new  branches  arise. 

Tomentose.  Covered  with  fine  “hair”;  having  a  downy  or  woolly  appear¬ 
ance. 

Trebouxia.  A  genus  of  single-celled  green  algae.  Its  distinctive,  single, 
disk-shaped  chloroplast  almost  fills  the  cell,  and  has  a  lobed  or 
crenate  margin.  It  is  the  most  common  green  phycobiont  in  lichens 
( figure  89B ) 

Trebouxioid.  Appearing  similar  to  Trebouxia. 

Trentepohlia.  A  genus  of  filamentous  green  algae  found  in  many  crustose 
lichens;  when  lichenized,  the  alga  often  produces  only  very  short 
filaments  or  is  single-celled.  The  orange-red  pigmented  globules, 
common  in  the  cells  of  unlichenized  individuals,  are  more  infrequent 
or  absent  in  lichenized  individuals  (figure  89A) 

Trichotomy.  See  branching. 

Truncate.  More  or  less  square  or  blunt  at  the  base. 

Tubercle.  A  minute,  wart-like,  thalline  protuberance  in  which  the  cortex 
is  generally  broken  at  the  apex. 

Umbilicus.  A  solitary,  short,  thick,  stem-like,  purely  hyphal  attachment 
organ  present  on  various  foliose  and  subfo'iose  lichens,  especially 
species  of  Umbilicaria. 

Uniseriate.  Spores  occurring  in  one  row  within  the  ascus. 

Vein.  In  lichens,  broad  or  narrow  ridges  or  thickenings,  often  pigmented, 
on  the  lower  surface  of  some  species  of  Peltigera. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  295 

Vermiform.  Shaped  like  a  worm:  i.e.,  elongate,  curved,  more  or  less 

rounded. 

Verruca  ( verrucae ).  A  conspicuous  wart-like  thalline  protuberance. 
Verruculose.  Covered  with  minute  verrucae. 

II.  CHEMICAL  TERMS 

All  lichen  substances  mentioned  in  the  keys  or  discussions  are 
listed  here  together  with  their  reactions  with  standard  color  test  re¬ 
agents  and  notes  on  their  identification  using  recrystallization  techniques. 

By  way  of  introduction  to  this  portion  of  the  glossary,  a  few  com¬ 
ments  on  general  methods  for  the  color  “spot”  tests  and  microchemical 
crystallization  are  presented.  Although  chromatography  was  used  exten¬ 
sively  in  some  parts  of  the  study,  the  techniques  and  data  are  too  extensive 
to  be  presented  here.  For  this  information,  Imshaug  and  Brodo  (1966) 
or  Hale  (1961a)  should  be  consulted. 

1.  Color  tests.  Reagents  (KOH,  Chlorox,  iodine)  should  be  stored 
in  small  jars  or  bott'es.  Since  alcoholic  solutions  of  PD  are  very  unstable, 
and  soon  after  preparation  are  unusable,  small  quantities  of  fresh  PD 
should  be  prepared  as  needed  (see  glossary  entry  under  PD). 

All  reagents  should  be  applied  to  the  thalli  using  a  capillary  pipette 
(such  as  a  melting  point  tube)  and  never  with  the  dropper  from  a 
reagent  bottle.  The  pipettes  can  be  drawn  to  a  fine  point  for  even  better 
control  of  the  reagent.  Allow  the  reagent  to  pass  into  the  pipette  by  capil¬ 
lary  action,  and  merely  touch  the  tube  to  the  lichen  material  to  empty  a 
tiny  but  adequate  amount  on  the  area  to  be  tested.  Results  should  be 
observed  under  a  dissecting  microscope.  KOH  and  PD  colors  are  perma¬ 
nent  and  will  often  darken  with  time,  but  C  and  KC  reactions  are  tem¬ 
porary  and  ephemeral. 

For  medullary  reaction  tests,  expose  a  small  portion  of  the  medulla 
by  cutting  away  the  cortex  with  a  razor  blade.  Reagents  may  be  applied 
to  any  undamaged  portion  of  the  cortex  for  cortical  tests.  Tested  portions 
of  the  thalli  should  always  be  discarded. 

2.  Crystal  tests.  Many  lichen  substances  can  be  extracted  from 

the  intact  lichen  thallus  (or  apothecium)  and  recrystallized  into  a  char¬ 
acteristic  and  recognizable  form.  The  recrystallization  reagents  are  gen¬ 
erally  one  of  the  following:  G.E.,  G.A.W.,  G.A.oT.,  G.A.An., 

G.W.Py  (see  glossary  below  for  preparation  formu’as). 

An  extraction  is  made  as  follows:  A  small  portion  of  the  thallus 
or  a  few  apothecia  are  placed  in  the  center  of  a  perfectly  clean  micro¬ 
scope  slide  which  is  placed  on  a  slide  warming  table  set  at  60°C.  Acetone 
is  deposited  on  the  lichen  material  drop  by  drop  (allowing  each  drop 
to  evaporate  before  applying  the  next)  until  5  to  10  drops  have  been 
added.  Lichen  substances,  if  present,  will  appear  as  a  residue  ring 
around  the  lichen  material.  An  alcohol  lamp  or  microflame  bunsen  burner 
can  be  used  instead  of  a  slide  warming  table,  but  open  flames  should 
be  used  with  caution  because  of  the  inflammability  of  acetone. 


296  THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 

The  lichen  material  is  now  discarded  (or,  if  scarce,  saved  for 
morphological  studies).  The  residue  is  generally  scraped  together,  using 
a  clean  razor  blade.  A  small  drop  of  the  proper  reagent  is  placed  on  the 
residue  and  a  clean  cover  glass  carefully  lowered  into  place.  The  slide  is 
once  again  warmed  for  about  one  minute  on  the  warming  table.  If  a  flame 
is  used,  special  care  must  be  taken  so  as  to  prevent  the  material  from 
boiling.  The  slide  is  then  allowed  to  cool. 

Some  crystals  appear  almost  immediately  (e.g.,  atranorin),  and 
some  take  much  longer  (e.g.,  salacinic  acid).  Because  all  the  reagents  are 
made  with  glycerine,  the  slides  may  be  left  overnight  or  longer,  if 
necessary,  and  they  will  not  dry  out.  Crystals  should  be  observed  with  a 
compound  microscope. 

Alectoronic  acid.  An  orsellic  acid  depsidone;  PD  — ,  KOH  — ,  KC  +  red, 
C  — ;  in  G.A.W. :  colorless,  radiating,  irregular  lamellae. 
Anthraquinone.  A  bright  red,  orange,  or  yellow  pigment  found  in  many 
lichenized  and  some  unlichenized  fungi;  turns  a  deep  red  or  purple 
upon  application  of  KOH. 

Atranorin.  A  /J-orsellic  acid  depside  found  in  many  lichens;  PD  —  or  + 
faint  yellow  (depending  on  concentration),  KOH  +  yellow,  KC  — , 
C  — ;  in  G.A.oT.  solution:  yellow,  straight  or  curved,  usually  highly 
branched,  very  slender  needles;  in  G.E.:  colorless,  straight,  blade¬ 
shaped  crystals. 

Baeomycic  acid.  A  /Torsellic  acid  depside;  PD  +  lemon  yellow,  KOH  — , 
KC  — ,  C— ;  in  G.A.  An.:  yellow,  thick  needles  often  with  frayed 
ends,  often  slow  in  forming. 

Barbaric  acid.  A  /J-orsellic  acid  depside;  PD  — ,  KOH  — ,  KC  +  orange, 
C  — ;  in  G.E.  solution:  colorless,  short,  prismatic  crystals;  in 
G.W.Py. :  colorless,  narrow,  rectangular  lamellae,  often  appearing 
as  if  the  ends  are  broken  off. 

Barbatolic  acid.  A  rare  lichen  substance;  PD  +  yellow,  KOH  +  yellow, 
KC  +  red,  C 

C.  Undiluted  household  bleach  (sodium  hypochlorite  solution);  deterio¬ 
rates  rapidly  and  therefore  must  be  poured  fresh  every  few  days. 

C operatic  acid.  A  fatty  acid;  PD  — ,  KOH  — ,  KC  — ,  C  — ;  in  G.E.: 

irregular,  “warty,”  subglobular  clumps  of  colorless  crystals. 
Cryptochloropliaeic  acid.  A  lichen  acid;  PD  — ,  KOH  — ,  KC  +  red, 
C  — ;  in  G.A.W. :  colorless,  extremely  slender,  abundantly  branched, 
curved  or  curled  needles. 

Didymic  acid.  A  dibenzofurane  compound  known  from  several  species  of 
Cladonia;  PD  -,  KOH  -,  KC  -,  K  -;  in  G.A.W.:  colorless, 
slender  needles,  slightly  or  strongly  curled  or  hooked  at  the  ends; 

in  small  clusters. 

Divaricatic  acid.  An  orsellic  acid  depside;  PD  — ,  KOH  — ,  KC  — ,  C  — ; 
in  G.E.  or  G.A.W.:  colorless  or  pale  yellow  straight  or  slightly 
curved  needles,  producing  conspicuous  perpendicular  branches;  often 
in  radiate  clusters. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


297 


Fumarprotocetraric  acid.  A  /3-orsellic  acid  depsidone;  PD+  red,  KOH  — 
(or  +  dingy  brown),  KC  — ,  C  — ;  cannot  be  dependably  demon¬ 
strated  by  crystal  tests. 

G.A.An.  Glycerin  —  95  percent  ethanol  • — ■  aniline,  2:2:1 

G.A.oT.  Glycerin  - —  95  percent  ethanol  —  o-toluidine,  2:2:1 

G.A.W.  Glycerin  —  95  percent  ethanol  —  water,  1:1:1 

G.E.  Glycerin  —  glacial  acetic  acid,  1:1 

Grayanic  acid.  A  lichen  acid  found  in  a  few  species  of  Cladonia ;  PD  — , 
KOH~,  KC  — ,  C— ;  in  untreated  acetone  extract  residue:  colorless, 
very  long,  straight  needles,  sometimes  becoming  blade-shaped;  in 
G.A.W. :  colorless,  slender,  straight  and  unbranched  needles  some¬ 
times  occurring  in  clusters. 

G.W.Py.  Glycerin  —  water  • — -  pyridine,  1:3:1 

Gyrophoric  acid.  An  orsellic  acid  depside;  PD  — ,  KOH  —  ,  KC+  red, 
C+  red;  in  G.A.W.  solution:  colorless,  small,  granule-like  clusters 
of  crystals. 

Homosekikaic  acid.  An  orsellic  acid  depside  found  only  in  Cladonia 
nemoxyna  (on  Long  Island):  PD  —  ,  KOH  —  ,  KC  — ,  C— ;  in  G.A.oT. 
solution  (after  scraping  acetone  extract  together  and  applying  the 
solution  to  the  underside  of  a  cover  slip):  oily  yellow  masses  in 
which  are  formed  yellow,  irregular,  very  thin  lamellae. 

I.  A  1  percent  solution  of  iodine  in  10  percent  potassium  iodide,  or  an 
alcoholic  solution  made  by  dissolving  a  few  crystals  of  iodine  in 
10  ml  of  70  percent  ethanol.  Iodine  tests  should  normally  be  done 
with  sectioned  material  under  magnification. 

Imbricaric  acid.  An  orsellic  acid  depside.  PD  —  ,  KOH  —  ,  KC  — ,  C  — ; 
microchemical  methods  cannot  distinguish  this  substance  from  simi¬ 
lar  perlatolic  acid  (Culberson,  1958b). 

KC.  A  reagent  combination  used  in  color  tests.  The  area  to  be  tested 
is  moistened  with  KOH,  after  which  C  is  applied.  A  positive  re¬ 
action  (usually  a  rose  or  orange  color)  is  usually  very  fleeting,  and 
must  be  observed  carefully  under  magnification. 

KOH  (K).  A  10-20  percent  solution  of  potassium  hydroxide. 

KOH  +  K2C03  (K~).  5  percent  KOH  —  20  percent  K2C03,  1:1 

Lobaric  acid.  An  orsellic  acid  depsidone;  PD  —  ,  KOH  —  ,  KC+  red,  C— ; 
in  G.A.W.:  colorless  crystals,  fanning  out  in  curved  radiate  clusters; 
difficult  to  distinguish  from  some  other  substances,  especially 
lecanoric  acid. 

Merochlorophaeic  acid.  A  rare  lichen  acid:  PD  — ,  KOH+  wine  red  (?), 
KC  — ,  C— ;  in  G.E.  solution:  colorless,  narrow  lamellae  with  oblique 
ends,  radiating  out  from  a  common  point. 

Monoacetyl-protocetraric  acid.  A  ^-orsellic  acid  depsidone:  PD+  red- 
orange,  KOH  —  ,  KC  ?,  C  ?.  The  crystal  forms  in  G.E.  (colorless, 
flat,  and  blade-like)  are  difficult  to  distinguish  from  crystals  pro¬ 
duced  by  atranorin.  It  is  best  identified  using  chromatography 
(Imshaug  and  Brodo,  1966). 


298  THE  UN.VERSITY  CF  THE  STATE  OF  NEW  YORK 

Norstictic  acid.  A  /j-orsellic  acid  depsidone;  PD  +  yellow,  KOH  + 
yellow  becoming  blood  red,  KC  — ,  C  — ;  in  KOH  or  KOH  + 
KjCO^:  orange  or  red,  short,  acicular  crystals,  clustered  or  solitary; 
in  G.A.oT.:  yellow,  very  thin,  square  or  rectangular  or  sometimes 
irregular  lamellae,  often  overlapping  in  small  clusters. 

Olivetoric  acid.  An  orsellic  acid  depside;  PD  — ,  KOH  — ,  KC  +  red, 
C  +  red;  in  G.A.W.:  colorless,  long,  very  slender,  curved  needles. 

Parietin.  A  yellow  or  orange  anthraquinone  pigment  commonly  found  in 
members  of  the  Teloschistaceae. 

PD.  A  freshly  prepared,  very  dilute  solution  of  para-phenylenediamine 
in  95  percent  ethanol.  It  is  best  prepared  on  a  glass  depression 
microscope  slide  by  adding  a  drop  or  two  of  the  alcohol  to  a  very 
small  quantity  of  the  chemical  (enough  to  cover  the  tip  of  a  dis¬ 
secting  needle).  For  larger  quantities  of  PD,  equivalent  proportions 
of  the  reagents  should  be  used.  The  material  is  extremely  toxic  and 
can  easily  stain  the  table  surface,  clothing,  and  herbarium  packets 
and  so  should  be  handled  and  applied  with  care. 

Perlatolic  acid.  An  orsellic  acid  depside;  PD  —  ,  KOH  —  ,  KC  — ,  C— ; 
in  G.A.W.  (after  concentrating  the  acetone  extract  residue):  color¬ 
less,  branched,  slightly  curved  or  straight,  long  needles. 

Physodic  acid.  An  orsellic  acid  depsidone;  PD  — ,  KOH  — ,  KC  +  red, 
C  — ;  in  G.A.W. :  colorless,  short,  curved  and  branching  crystals. 

Protocetraric  acid.  A  /i-orsellic  acid  depsidone;  PD  +  red-orange, 
KOH  — ,  KC  +  red,  C  — ;  in  G.A.oT.:  yellow,  irregular,  granule¬ 
like  crystals. 

Protolichesterinic  acid.  A  lactonic  acid;  PD—,  KOH  — ,  KC  — ,  C— ; 
in  G.E. :  color'ess,  square  or  rectangular,  thin  lamellae;  best  seen 
in  polarized  light. 

Pseudonorangiformic  acid.  A  lichen  acid  found  only  in  Cladonia  submitis; 
PD  — ,  KOH  — ,  KC  — ,  C  — ;  in  G.E.:  colorless  crystals,  falcate  or 
arborescent,  or  in  circular,  curled  clusters;  crystalizes  very  slowly. 

Psoromic  acid.  A  /^-orsellic  acid  depsidone;  PD  +  deep  yellow,  KOH  — , 
KC  — ,  C— ;  in  G.E.:  colorless,  feather-like  fascicles  of  slender 
curved  needles. 

Pnlvic  acid  derivative.  A  yellow,  KOH  —  pigment  such  as  is  found  in 
Candelaria. 

Salacinic  acid.  A  /^-orsellic  acid  depsidone;  PD  +  yellow,  KOH  +  yellow 
slowly  turning  blood  red,  KC  — ,  C  — ;  in  KOH  +  KL.CO:!:  dark 
red  curved  needles  in  tightly-bound  fascicles  recembling  sheaves  of 
wheat;  often  very  slow  in  forming,  especially  when  the  concentra¬ 
tion  is  low;  in  G.A.oT.:  yellow,  small,  boat-shaped  (fusiform) 
crystals,  often  in  small  clusters. 

Squamatic  acid.  A  /^-orsellic  acid  depside;  PD  — ,  KOH  — ,  KC  — ,  C  — ; 
bright7 y  fluorescent  (blue-white)  in  ultraviolet  light;  in  G.E.:  color¬ 
less,  short  prisms,  resembling  rice  grains,  sometimes  in  small  clusters, 
but  usually  solitary. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK  299 

Stictic  acid.  A  /J-orsellic  acid  depsidone;  often  found  in  conjunction  with 
norstictic  acid,  either  in  the  same  thallus,  or,  in  a  corresponding 
and  closely  related  species;  PD  +  pale  orange,  KOH  +  deep  yellow, 
KC  — ,  C  — ;  in  G.A.oT:  very  pale  yellow,  small,  thin,  hexagonal 
lamellae. 

Strepsilin.  A  dibenzofurane  present  only  in  Cladonia  strepsili$\  PD  — , 
KOH  — ,  KC  +  green,  C  +  green. 

Substance  H.  A  lichen  substance  found  in  Cladonia  conista;  PD  — , 
KOH  — ,  KC  — ,  C  — ;  in  an  untreated  acetone  extract  allowed  to 
dry  on  the  slide:  long,  colorless  needles,  parallel  at  the  center  but 
irregularly  radiating  at  the  periphery  of  the  residue. 

Thamnolic  acid.  A  /3-orsellic  acid  depside;  PD  +  orange,  KOH  +  deep 
yellow,  KC  — ,  C  — ;  in  G.A.An.:  yellowish,  straight,  slender  needles 
grouped  into  fascicles  like  sheaves  of  wheat. 

Usnic  acid.  A  yellow  dibenzofurane  pigment;  one  of  the  most  common 
lichen  substances;  PD  — ,  KOH  — ,  KC  +  yellow  or  orange  (faint), 
C  — ;  in  G.E.:  yellow,  narrow,  flat  needles,  sometimes  broadening 
into  lamellae,  often  clustered. 

UV.  Ultraviolet  light. 

Variolaric  acid.  A  lichen  substance  found  in  some  species  of  Ochrolechia 
(diagnostic  test  on  p.  221). 

Zeorin.  A  triterpenoid  substance  (aliphatic);  PD  — ,  KOH  — ,  KC  — , 
C— ;  in  G.A.An.  or  G.A.oT.:  colorless,  double  pyramid  crystals, 
often  showing  a  conspicuous  equatorial  expansion. 


Appendix  C 

CHECKLIST  OF  THE  LICHENS  OF  LONG  ISLAND 


Class  ASCOMYCETES 
Subclass 

LOCULOASCOMYCETIDAE 

Order  Pleosporales 
Arthopyreniaceae 

Arthopyrenia  cerasi  (Schrad.)  Mass. 

A.  pinicola  (Hepp)  Mass. 

Leptorhaphis  epidermidis 

(Ach.)  Th.  Fr. 

Polyblastiopsis  quercicola  Brodo 

Order  Myrangiales 
Arthoniaceae 

Arthonia  caesict  (Flot.)  Korb. 

A.  mediella  Nyl. 

A.  polymorpha  Ach. 

A.  punctiformis  Ach. 

A .  sexlocularis  Zahlbr. 

A.  siderea  Degel. 

A  rthotheUum  taediosum 
(Nyl.)  Miill.  Arg. 

Micarea  melaena  (Nyl.)  Hedl. 

M.  prasina  (Fr.)  Korb. 

Order  Hysteriales 
Opegraphaceae 
Opegrapha  cinerea  Chev. 

O.  rufescens  Pers. 

Subclass  ASCOMYCETIDAE 

Order  Caliciales 
Caliciaceae 

Chaenotheca  phaeocephala 

(Turn.)  Th.  Fr. 

Order  Sphaeriales 
Verrucariaceae 
Verrucaria  microspora  Nyl. 

V .  muralis  Ach. 

V .  nigrescens  Pers. 

V.  silicicola  Fink  in  Hedr. 
Dermatocarpon  miniatum  (L.)  Mann 
Pyrenulaceae 

Pyrenida  nitida  (Weig.)  Ach. 
Melanotheca  cruenta  (Mont.) 

Mull.  Arg. 

Trypethelium  virens  Tuck,  in  W.  Dari. 
Porinaceae 


Porina  cestrensis  (Tuck,  in  W.  Dari.) 
Mull.  Arg. 

P.  hibernica  P.  James  &  Swins. 

in  Swins. 

P.  nucula  Ach. 

Order  Lecanorales 
Graphidaceae 

Xylographa  opegraphella  Will, 
in  Rothr. 

Grapliis  scripta  (L.)  Ach. 
Phaeograpliis  dendritica  (Ach.) 

Mull.  Arg. 

Diploschistaceae 

Diploschistes  scruposus  (Schreb.) 
Norm. 

Gyalectaceae 

Dimerella  diluta  (Pers.)  Trev. 

D.  lutea  (Dicks.)  Trev. 
Collemataceae 

Collema  subfurvum  (Miill.  Arg.) 
Degel. 

Leptogium  corticola  (Tayl.)  Tuck, 
in  Lea 

L.  cyanescens  (Ach.)  Korb. 
Pannariaceae 

Placynthium  nigrum  (Finds.)  S.  Gray 
P armaria  lurida  (Mont.)  Nyl. 
Stictaceae 

Lobaria  pulmonaria  (L.)  Hoffm. 

L.  quercizans  Michx. 

Nephromaceae 
Nephroma  laevigatum  Ach. 
Peltigeraceae 

Solorina  saccata  (L.)  Ach. 

Pcltigera  aphthosa  (L.)  Willd. 

P.  canina  (L.)  Willd. 

P.  polydactyla  (Neck.)  Eloffm. 

P.  praetextata  (Florke  in  Somm.) 
Vain. 

Lecideaceae 

Lecidea  aeruginosa  Borr.  in 
Hook.  &  Sowerby 
L.  albocaerulescens  (Wulf.  in 
Jacq.)  Ach. 

L.  anthracophila  Nyl. 

L.  botryosa  (Fr.)  Th.  Fr. 


300 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


301 


L.  coarctata  (Turn,  in  Sm.  & 

Sowerby)  Nyl. 

L.  cyrtidia  Tuck. 

L.  erratica  Kerb. 

L.  granulosa  (Ehrh.)  Ach. 

L.  macrocarpa  (DC.  in  Lam.  &  DC.) 
Steud. 

L.  myriocarpoides  Nyl. 

L.  nylanderi  (Anzi)  Th.  Fr. 

L.  scalaris  (Ach.)  Ach. 

L.  uliginosa  (Schrad.)  Ach. 

L.  various  Ach. 

L.  vernalis  (L.)  Ach. 

L.  viridescens  (Schrad.  in  Gmel.) 

Ach. 

Catillaria  glauconigrans  (Tuck.)  Hasse 
Bacidia  atrogrisea  (Del.  in  Hepp) 

Korb. 

B.  chlorantha  (Tuck.)  Fink 
B.  chlorococca  (Graewe  in  Stizenb.) 
Lett. 

B.  chlorosticta  (Tuck.)  Schneid. 

B.  intermedia  (Hepp  in  Stizenb.)  Arn. 
B.  inundata  (Fr. )  Korb. 

B.  schweinitzii  (Tuck,  in  W.  Dari.) 
Schneid. 

B.  trisepta  (Naegeli  in  Mull.  Arg.) 
Zahlbr. 

B.  umbrina  (Ach.)  Bausch. 
Rliizocarpon  einereovirens  (Miill. 

Arg.)  Vain. 

R.  grande  (Florke  in  Flot.)  Arn. 

R.  intermedium  Degel. 

R.  obscuration  (Ach.)  Mass. 

R.  plicatile  (Leight.)  A.L.  Sm. 
Stereocaulaceae 

Pycnothelia  papillaria  (Ehrh.)  Duf. 
Stereocaulon  saxatile  Magn. 

Baeomycetaceae 
Baeomyces  roseus  Pers. 

Cladoniaceae 

Cladonia  alpestris  (L.)  Rabenh. 

C.  apodocarpa  Robb. 

C.  arbuscula  (Wallr.)  Rabenh. 

C.  atlantica  Evans 
C.  bacillaris  (Ach.)  Nyl. 

C.  beaumontii  (Tuck.)  Vain. 

C.  boryi  Tuck. 

C.  brevis  Sandst. 

C.  caespiticia  (Pers.)  Florke 
C.  calycantha  Nyl. 


C.  capitata  (Michx.)  Spreng. 

C.  carassensis  Vain. 

C.  cariosa  (Ach.)  Spreng. 

C.  carneola  (Fr.)  Fr. 

C .  caroliniana  Tuck. 

C.  chiorophaea  (Florke  in  Somm.) 
Spreng. 

C.  ciavulijera  Vain. 

C.  coniocraea  (Florke)  Spreng.  em. 
Sandst. 

C.  conista  (Ach.)  Robb. 

C.  cristatella  Tuck. 

C.  cylindrica  (Evans)  Evans 
C.  deformis  (L.)  HofTm. 

C.  didyma  (Fee)  Vain. 

C.  evansii  Abb. 

C.  farinacea  (Vain.)  Evans 
C.  fimbriata  (L.)  Fr. 

C.  floerkeana  (Fr.)  Florke 
C.  floridana  Vain. 

C.  furcata  (Huds.)  Schrad. 

C.  incrassata  Florke 
C.  macilenta  HofFm. 

C.  mateocyatha  Robb. 

C.  mitis  Sandst. 

C.  multiformis  Merr. 

C.  nemoxyna  (Ach.)  Arn. 

C.  parasitica  (Hoffm.)  Hoffm. 

C.  piedmontensis  Merr. 

C.  pityrea  (Florke)  Fr. 

C.  pleurota  (Florke)  Schaer. 

C.  pyxidata  (L.)  Hoffm. 

C.  rangiferina  (L.)  G.Web.  in  Wigg. 

C.  robbinsii  Evans 
C.  santensis  Tuck. 

C.  scabriuscula  (Del.  in  Duby)  Nyl. 

C.  simulata  Robb. 

C.  squamosa  (Scop.)  Hoffm. 

C.  strepsilis  (Ach.)  Vain. 

C.  subcariosa  Nyl. 

C.  submitis  Evans 
C.  subtenuis  (Abb.)  Evans 
C.  terrae-novae  Ahti 
C.  uncialis  (L.)  G.Web. 

C.  verticillata  (Hoffm.)  Schaer. 

C.  vulcanica  Zoll. 

Umbilicariaceae 

Umbilicaria  mammulata  (Ach.)  Tuck. 
U.  muhlenbergii  (Ach.)  Tuck. 

U.  papulosa  (Ach.)  Nyl. 
Acarosporaceae 


302 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


Sarcogyne  clavus  (Ram.  in  Lam. 

&  DC. )  Kxemp. 

S.  privigna  (Ach.)  Mass. 

S.  simplex  (Dav.)  Nyl. 

Acarospora  juscata  (Schrad.)  Arn. 

Pertusariaceae 
Pertusaria  alpina  Hepp 
P.  amara  (Ach.)  Nyl. 

P.  multipuncta  (Turn.)  Nyl. 

P.  propinqua  Mull.  Arg. 

P.  subpertusa  Brodo 
P.  trachythallina  Erichs,  in  Degel. 

P.  tuberculifera  Nyl. 

P.  velata  (Turn.)  Nyl. 

P.  xanthodes  Mull.  Arg. 

Melanaria  macounii  Lamb 
Lecanoraceae 

Ionaspis  odorn  (Ach.  in  Schaer. ) 

Th.  Fr. 

Lecanora  atra  (Huds.)  Ach. 

L.  caesiocinerea  Nyl. 

L.  caesiorubella  Ach. 

L.  chlaroteru  Nyl. 

L.  cinerea  (  L. )  Somm. 

L.  coiiizaea  (Ach.)  Nyl. 

L.  cupressi  Tuck. 

L.  degelii  Schauer  &  Brodo 
L.  disperse!  (Pers.) 

L.  Iiageni  (Ach.)  Ach. 

L.  I cievis  Poelt 

L.  muralis  (Schreb.)  Rabenh. 

L.  rubina  (Vill.)  Ach. 

L.  subintricata  (Nyl.)  Th.  Fr. 

L.  symmicta  (Ach.)  Ach. 

L.  cfr.  varia  (Ehrh.)  Ach. 

L.  sp. 

Ochrolechia  parella  (L. )  Mass. 

O.  rosella  (Miill.  Arg.)  Vers. 

O.  sp. 

Haematomma  ochrophaeum  (Tuck.) 
Mass. 

//.  sp. 

Candelariaceae 

Candelariella  aurella  (Hoffm.) 
Zahlbr. 

C.  vitelline i  (Ehrh.)  Miill.  Arg. 
Candelaria  concolor  (Dicks.) 

B. Stein  in  Cohn 
Parmeliaceae 

Parmeliopsis  aleurites  (Ach.)  Nyl. 

P.  ambigua  (Wulf.  in  Jacq.)  Nyl. 


P.  placorodia  (Ach.)  Nyl. 

Parmelia  appalachensis  W.  Culb. 

P.  arseneana  Gyeln. 

P.  aurulenta  Tuck. 

P.  caperata  (L.)  Ach. 

P.  conspersa  (Ach.)  Ach. 

P.  dilatata  Vain. 

P.  galbina  Ach. 

P.  hypotropa  Nyl. 

P.  livida  Tayl. 

P.  michauxiana  Zahlbr. 

P.  olivetorum  Nyl. 

P.  perforata  (Wulf.  in  Jacq.)  Ach. 

P.  perlata  (Huds.)  Ach. 

P.  plittii  Gyeln. 

P.  reticulata  Tayl.  in  Mack. 

P.  rudecta  Ach. 

P.  saxatilis  (L.)  Ach. 

P.  stenophyUa  (Ach.)  Heug. 

P .  subaurifera  Nyl. 

P.  subrudecta  Nyl. 

P.  sulcata  Tayl.  in  Mack. 

P.  tasmanica  Hook.  &  Tayl. 
Hypogymnia  physodes  (  L. )  Nyl. 
Pseudevernia  furfuracea  ( L. )  Zopf 
Cetraria  ciliaris  Ach. 

C.  fendleri  (Nyl.)  Tuck. 

C.  islandica  (L. )  Ach. 

C.  tuckermanii  Oakes  in  Tuck. 

C.  viridis  Schwein. 

Anzia  colpodes  (Ach.)  Stizenb. 
Usneaceae 

Evernia  mesomorpha  Nyl. 

Alectoria  glabra  Mot. 

A.  nidulifera  Norrl.  in  Nyl. 

Ramalina  complanata  (Sw.  in  Ach.) 

Ach. 

K.  fastigiata  (Lilj.)  Ach. 

R.  stenospora  Miill.  Arg. 

R.  willeyi  R.H.  Howe 
Us nea  longissima  Ach. 

U.  mutabilis  Stirt. 

U.  strigosa  (Ach.)  A.  Eaton 
U.  tricliodea  Ach. 

U.  sp.  Teloschistaceae 

Caloplaca  aurantiaca  (Lightf. )  Th.  Fr. 

C.  camptidici  (Tuck.)  Zahlbr. 

C.  cerina  (Ehrh.  in  HofFm.)  Th.  Fr. 

C.  citrina  (Hoffm.)  Th.  Fr. 

C.  discolor  (Will,  in  Tuck.)  Fink 
C.  feracissima  Magn. 


LICHENS  OF  LONG  ISLAND,  NEW  YORK 


303 


C.  flavovirescens  (Wulf.)  Dalla 
Torre  &  Sarnth. 

C.  pyracea  (Ach.)  Th.  Fr. 
Xanthoria  falla.x  (Hepp  in  Arn.) 
Arn. 

X.  parietina  (L.)  Beltr. 
Teloschistes  chrysophthalmus 

(L.)  Beltr. 

T.  flavicans  (Sw. )  Norm. 
Physciaceae 

Buellia  curtisii  (Tuck.)  Irtish, 
in  Brodo 

B.  dialyta  (Nyl.)  Tuck. 

B.  polvspora  (Will,  in  Tuck.) 
Vain. 

B.  punctata  (HofFm.)  Mass. 

B.  stigmaea  Tuck. 

B.  stillingiana  J.  Stein 
B.  turgescens  Tuck. 

Rinodina  applanata  Magn. 

R.  confragosa  (Ach.)  Korh. 

R.  milliaria  Tuck. 


R.  oreina  (Ach.)  Mass. 

R.  pachysperma  Magn. 

R.  salina  Degel. 

Pyxine  sorediata  (Ach.) 

Mont,  in  Sagra 

Physcia  adscendens  (Th.  Fr.)  Oliv. 
P.  aipolia  (Ehrh.  in  Humb.) 

Hampe  in  Fiirnr. 

P.  millegrana  Degel. 

P.  orbicularis  (Neck.)  Potsch  in 
Potsch  &  Scheiderm. 

P.  stellaris  (L. )  Nyl. 

P.  subtilis  Degel. 

P.  tribacoides  Nyl. 

Anaptychia  obscurata  (Nyl.)  Vain. 
A.  palmulata  (Michx. )  Vain. 

A.  pseudospeciosa  Kurok. 

Class  FUNGI  IMPERFECTI 
Lepraria  incana  (  L. )  Ach. 

L.  zonal  a  Brodo 
L.  sp. 


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Hale,  M.  E.,  Jr.  1950.  The  lichens  of  Aton  Forest,  Connecticut,  Bryologist  53: 
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Johnson,  G.  T.  1940.  Contributions  to  the  study  of  the  Trypetheliaceae.  Ann.  Mo. 
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Kuchler,  A.  W.  1964.  Potential  natural  vegetation  of  the  conterminous  United 
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Mozingo,  H.  N.  1961.  The  genus  Cladonia  in  eastern  Tennessee  and  the  Great 
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314 


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LICHENS  OF  LONG  ISLAND,  NEW  YORK 


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Index  to  Names 


Names  in  italics  are  synonyms;  numbers  in  italics  indicate  a  position  in  the 
key;  numbers  in  boldface  indicate  a  main  heading  in  the  annotated  list;  an 
asterisk  (*)  next  to  a  number  indicates  a  figure. 


Abies  217 

abietina  (Xylographa)  158 
Acarospora  205 
Acarosporaceae  204 

Acer  33 

acerifolium  (Viburnum)  13 
Actinogyra  204 
aculeata  (Cornicularia)  60 
adscendens  (Physcia)  30.  81,  145, 
258,  259 

adscendens  (Physcia  stellaris  var.)  258 
aeruginosa  (Crocynia)  263 
aeruginosa  (Lecidea)  22.  56,  81,  118 , 
126,  165,  167,  169,  220,  273.  281 
aipolia  (Physical  81,  96*,  145.  259, 
267,  271.  274 
aipolius  (Lichen)  259 
alba  (Hicoria)  9 

alba  (Quercus)  9.  12.  13,  14*,  21,  25, 
26,  27,  28,  37,  49,  50,  51.  52,  53, 
54,  146,  147,  148,  210,  221,  227, 
228.  262,  263 
albescens  (Bacidia)  174 
albescens  (Peltigera  canina  var.)  164 
albocaerulescens  (Lecidea)  55,  81, 
100*,  117,  125,  166,  271,  273 
albocaerulescens  (Lichen)  166 
Alectoria  122,  141,  241 
aleurites  (Lichen)  226 
aleurites  ( Parmeliopsis)  49,  56,  57, 
81,  139,  226,  274 
allophana  (Lecanora)  219 
alnifolia  (Clethra)  16 
Alpestres  (Cladonia  sect.)  199 
alpestris  (Cladonia)  68*,  80,  128,  199. 

201,  275,  276,  277 
alpestris  (Lichen  rangiferinus  [7])  199 
alpina  (Pertusaria)  136,  206 
amara  (Pertusaria)  57,  81,  94*,  7/7, 
1 36.  206.  237,  275 
amara  (Variolaria)  206 
ambigua  (Buellia)  258 


ambigua  (Parmeliopsis)  56,  57,  80, 
121,  139,  226 
atnbiguus  (Lichen)  226 
americana  (Alectoria)  241 
americana  (Ramalina)  243 
americana  (Ulmus)  25,  27,  49.  54, 
146 

Amphigymnia  ( Parmelia  subgenus) 
229,  231,  284 

Ampliaria  (Pertusaria  subgenus)  284 
amplissima  (Lobaria)  87,  88,  162 
Anaptychia  121 ,  146,  262 
Andropogon  42 
angustifolium  (Vaccinium)  12 
anthracophila  (Lecidea)  21,  45,  49, 
57,  81,  95*,  117,  125,  129,  166, 
170,  267,  271,  273 
anthracophila  (Psora)  166 
Anzia  240.  287 

aphthosa  (Peltigera)  80,  102*,  1 20, 
124,  163,  275 

aphthosa  (Peltigera  aphthosa  var.)  163 
aphtosus  (Lichen)  163 
apodocarpa  (Cladonia)  81,  129,  192 
appalachensis  (Parmelia)  70*,  82,  139, 

111 

applanata  (Rinodina)  145,  256 
aquila  (Anaptychia)  262 
aquilinum  (Pteridium)  12 

arbuscula  (Cladonia)  42,  80,  129, 

200.  201,  202,  203,  276 
arbuscula  (Cladonia  arbuscula  subsp.) 
202 

Arbuscula  (Patellaria  foliacea  var. 

m.)  201 

arceutina  (Bacidia)  174 

arizonica  (Usnea)  246 

arseneana  (Parmelia)  29,  55,  139, 

111 

Arthoniu  113,  122,  150.  286 
Arthoniaceae  150 
Arthopyrenia  114,  122,  146 
Arthopyreniaceae  146 


318 


INDEX  TO  NAMES 


319 


Arthothelium  151 

Asccmycetidae  284 
Aspicilia  (Lecanora  sect.)  136 ,  216, 
285 

assericola  (Candelariella  vitellina  var.) 
225 

ater  (Lichen)  213 

atlantica  (Cladonia)  14*.  56,  57,  60, 
82,  109.  I  10.  130,  134,  193.  194, 
271,  272 

atra  (Lecanora)  81.  137,  213 
atrogrisea  (Bacidia)  81,  127 ,  172 
citrogrisea  (Biatora)  172 
attenuata  (Ramalina)  244 
aurantiaca  (Caloplaca)  81,  143,  248 
aurantiacus  (Lichen)  248 
aurella  (Candelariella)  21.  59,  80. 
138,  224 

aurella  (Candelariella  aurella  var.) 
225 

aurella  (Verrucaria)  224 
aurulenta  (Parmelia)  75*.  81,  140, 
111,  267,  275 

austroamericana  (Gchrolechia)  223 
austroamericana  ( Ochrolechia  palles- 
cens  var.)  223 

austroamericana  ( Ochrolechia  tarta- 
rea  var.)  223 

baccata  (Gaylussacia)  12 
Bacidia  115,  126.  172 
bacillaris  (  Baeomyces )  1 80 
bacillaris  (Cladonia)  44,  49,  55,  56. 

81.  130,  133,  180.  195,  272,  275 
Baeomyces  26.  179 
Baeomycetaceae  111.  179 
banksiana  (Pinus)  226 
barbata  (Usnea)  245 
beaumontii  (Cladonia)  57,  82,  94*, 
7 30.  132,  194,  276 

beaumontii  (Cladonia  santensis  f. ) 
194 

Betula  148 
Biatora  174 
Biatorella  205 

Bicornutae  (Cladina  sect.)  198 
Bilimbia  152 
Blastenia  250 
bolliana  (Parmelia)  227 
borreri  (Parmelia)  235 


boryi  (Cladonia)  10*.  22,  32,  42.  56, 
60,  82,  103*,  128,  197,  200,  239, 
268,  271,  272 
botryosa  (Biatora)  166 
botryosa  (Lecidea)  56,  81.  118,  125, 
165,  166,  167 

breviligulata  (Ammophila)  10*.  12,  42 
brevis  (Cladonia)  81,  130,  132,  184, 

185 

brevispora  (Bacidia  arceutina  f. )  174 
Buellia  115,  144,  253,  257 

caesia  (Arthonia)  1 22,  150,  272 
caesium  (Coniangium)  150 
caesiocinerea  (Lecanora)  136,  213, 
214,  216 

caesiorubella  (Lecanora)  21,  34.  35, 
37,  40,  109,  137,  190,  214,  273.  279 
(see  also  “lathamii” ) 
caespiticia  (Cladonia)  44,  55,  81.  99*, 
129,  130,  192.  268 
caespiticus  (Baeomyces)  192 
Caliciaceae  153 

Caloplaca  115.  143,  172.  218.  225. 
248,  258 

calycantha  (Cladonia)  57,  83,  97*, 
129,  134,  186,  271 

camptidia  (Caloplaca)  82,  101*,  143, 

248 

camptidia  (Lecanora)  248 

Candelaria  225,  298 
candelaria  (Xanthoria)  251 
Candelariaceae  111.  224 
Candelariella  115.  138,  224 
canina  (Peltigera)  80.  110.  1 24,  163. 
165 

canina  (Peltigera  canina  var.)  164 

caninus  (Lichen)  163 

caperata  (Parmelia)  21,  35.  50,  55.  57. 

60.  81.  139,  228.  273.  275 
caperatus  (Lichen)  228 
capitata  (Cladonia)  81,  131,  184.  270, 
272,  275 

capitatum  (Helopodiunr)  184 
carassensis  (Cladonia)  83.  134,  196 
cariosa  (Cladonia)  80,  132,  184 
cariosus  (Lichen)  184 
carneola  (Cenomyce)  182 
carneola  (Cladonia)  80,  134,  182,  183 
carolinensis  (Myrica)  12 


320 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


caroliniana  (Cladonia)  81,  128,  197, 
111 

caroliniana  (Umbilicaria)  204 
carpinea  (Lecanora)  88 
Carya  25,  54,  153.  248,  249 

Catillaria  171,  176 
Celtis  151 

cerasi  (Arthopyrenia)  722,  146 
cerasi  (Verrucaria)  146 
cerina  (Caloplaca)  30,  81,  143,  248, 
249,  251 

cerinus  (Lichen)  248 
cestrensis  (Porina)  82,  124,  156,  157 
cestrensis  (Verrucaria)  156 
Cetraria  121,  141,  238 
cetrarioides  (Parmelia)  232 
cetrata  (Parmelia)  232 
Chaenotheca  153.  288 
chalybeia  (Catillaria)  59 
chalybeiformis  (Alectoria)  242 
Chamaecyparis  9,  47,  106,  1 17,  227, 
240.  280 

Chasmariae  (Perviae  subsect.)  128, 

192 

chlarona  (Lecanora)  215 
chlarotera  (Lecanora)  35,  50.  54,  137, 
214,  215,  216,  218,  221,  273 
chlorantha  (Bacidia)  82,  119,  127, 
172 

clilorantlia  (Biatora)  172 
chlorococca  (Bacidia)  49,  50,  54,  56, 
57.  82,  118,  126,  172,  173.  269,  271, 
272 

chlorococca  (Lecidea)  173 
clilorophaea  (Cenomyce)  185 
chlorophaea  (Cladonia)  34,  35,  36. 
43,  44,  50,  55,  56,  60,  77,  80,  129, 
134,  182.  185,  188,  189,  190.  273, 
275,  279 

chlorostica  (Bacidia)  82.  126,  173 
chlorostica  (Lecidea)  173 
chrysophthalmos  (Lichen)  252 
chrysophthalmus  (Teloschistes)  83, 
144,  252.  253,  276,  277 
ciliaris  (Cetraria)  56,  57,  81,  93*. 

141,  238,  240,  267,  271,  272,  275 
ciliata  (Physcia)  261 
cinerea  (Cladonia  subtenuis  f. )  199, 
200 


cinerea  (Lecanora)  55,  80,  99*.  117, 
136,  166,  213,  216 

cinerea  (Opegrapha)  47,  100*,  119, 
123,  153 

cinereofusca  (Lecanora)  215 
cinereovirens  (Patellaria)  176 
cinereovirens  (Rhizocarpon)  127,  176, 
255 

cinereus  (Lichen)  216 
cinnabarina  (Arthonia)  151 
citrina  (Caloplaca)  21,  59,  81,  104*, 
116,  143,  249,  250.  269 
citrina  (Verrucaria)  249 
Cladina  (Cladonia  sect.)  201 
Cladina  (Cladonia  subgenus)  10*, 
111,  128,  198,  284,  285,  292 
Cladonia  2,  3,  34,  42,  43.  44,  107, 
111,  114,  117,  119,  121,  127,  159, 
178,  179,  285,  291,  292,  293,  296, 
297 

Cladonia  (Cladonia  subgenus)  179 
cladonia  (Parmelia)  87 
cladonia  ( Pseudevernia)  87,  88 
Cladoniaceae  179 
Clausae  (Cladonia  sect.)  179 
clavulifera  (Cladonia)  42,  56.  81,  129, 
131,  184,  185,  270,  271,  273 
davits  (Lichen)  204 
clavus  (Sarcogyne)  21,  55,  81,  135, 
204,  205 

coarctata  (Lecidea)  55,  81,  125,  167 
coarctatus  (Lichen)  167 
Cocciferae  (Clausae  subsect.)  179 
coccinea  (Quercus)  9,  12.  13,  14*, 
19,  21,  24,  26,  50,  51,  53,  54,  234 
coccinea  x  rubra  (Quercus)  19,  24, 
50,  51 

coccinea-velutina  (Quercus)  227 
Coccomyxa  120 

Coenogonium  108 
Collema  160 
Colleinntaceae  160 

colpodes  (Anzia)  71*,  81,  101*,  1 20, 
240,  241 

colpodes  (Lichen)  240 
comosa  (Usnea)  142 
complanata  (Ramalina)  83,  141,  242, 
244 

complanatus  (Lichen)  242 


INDEX  TO  NAMES 


321 


concolor  (Candelaria)  81,  117,  120, 
225,  267,  275 

concolor  (Candelaria  concolor  var.) 
225 

condensata  (Cladonia  impexa  f.)  198 
confragosa  (Parmelia)  256 
confragosa  (Rinodina)  81,  144,  256 
confusa  (Ramalina)  243 
coniocraea  (Cenomyce)  189 
coniocraea  (Cladonia)  35,  50,  60,  77, 
81,  129,  133,  188,  189,  190,  271, 
273 

coni st a  (Cenomyce  fimbriata  [3  C. ) 
188 

conista  (Cladonia)  81,  102*,  135, 

188,  189,  190,  275,  299 
conizaea  (Lecanora)  138,  217,  220, 
275 

conizaea  (Lecanora  expallens  fj  L.) 
217 

conradii  (Corema)  194 
conspersa  (Parmelia)  29,  55,  81.  139, 
176,  227.  228,  229,  233,  236,  273 
conspersus  ( Lichen  )  228 
contigua  (Lecidea)  169 
contigua  (Verrucaria)  169 
corticiia  (Collema)  160 
corticola  (Leptogium)  81.  124,  160 
corymbosum  ( Vaccinium)  14,  16,  25, 
50,  56.  227.  240.  267 
crispa  (Cetraria  islandica  subsp.)  32, 
56,  60,  1 10.  122,  141,  239,  271 
crispa  (Cetraria  islandica  7  C.)  239 
crispata  (Cladonia)  60 
cristatella  (Cladonia)  28,  43,  44,  49, 
55,  56.  60,  81.  110,  130,  131,  181, 
182,  190,  273,  275 
Crocynia  263 

cruenta  (Melanotheca)  82,  114,  155, 
156 

cruentum  (Trypethelium )  155 
cryptochlorophaea  (Cladonia)  188 
cupressi  (Lecanora)  82.  137,  217 
curtisii  (Buellia)  54,  59,  81.  118,  144, 
253,  254,  255.  272 
curtisii  (Gyrostomum)  253 
cyanescens  (Collema  tremelloides 
var.)  160 


cyanescens  (Leptogium)  81,  92*,  124, 
160,  161,  237,  269,  275,  291 
cyanescens  (Lichen)  160 
cyanescens  (Parmelia)  160 
cylindrica  (Cladonia)  83,  133,  190, 
191 

cylindrica  (Cladonia  borbonica  f. ) 
190 

cyrtidia  (Lecidea)  55,  81,  125,  167, 
168 

deformis  (Cladonia)  80,  134,  182 
deformis  (Lichen)  182 
degelii  (Lecanora)  137,  215,  217,  218, 
281 

delicata  (Cladonia)  192 
deltoides  (Populus)  55 
demissa  (Rinodina)  59,  258 
demissa  (Zeora  metabolica  ft)  258 
dendritica  (Opegrapha)  158 
dendritica  ( Phaeographis)  54,  59,  81, 
113,  158,  274 
dentata  (Castanea)  9 
dentatum  (Viburnum)  16 
Dermatocarpon  155 
dialyta  (Buellia)  82,  144,  253,  254 
dialyta  (Lecidea)  253 
didyma  (Cladonia)  57,  83,  94*,  109, 
130,  133,  181 

didymus  (Scyphophorus)  181 
digitata  (Cladonia)  134 
dilatata  (Parmelia)  83,  140,  229 
dillenii  (Gyrophora)  204 
dillenii  ( Umbilicaria)  204 
diluta  (Dimerella)  81,  116,  159 
diluta  (Peziza)  159 
Dimerella  159 
Diploscliistaceae  159 
Diplosehistes  159 

discolor  (Caloplaca)  115,  117,  143, 
249,  250 

discolor  (Placodium  ferrugineum 
var.)  249 

dispersa  (Lecanora)  21,  55.  59,  80. 

137,  205,  218 
dispersus  (Lichen)  218 
distendens  ( Arthothelium )  152 
Drosera  16 

dubia  (Opegrapha)  47 
dubia  (Parmelia)  235 


322 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


effusa  (Bacidia)  174,  175 
effusa  (Candelaria  concolor  var.)  225 
effusa  (Lecanora)  220 
effuse  (Theloschistes  concolor  var.) 
225 

effusus  (Lichen)  174 
elatinum  (Haematomma)  224 
endochrysea  (Physcia)  260.  261 
endochrysea  (Physcia  obscura  var.) 
260 

endococcinea  (Physcia)  260,  261 
endococcinea  (Physcia  abscura  f. ) 
260 

endoleuca  (Bacidia)  172 
epiclada  (Parmelia)  231 
epidermidis  (Leptorhaphis)  21,  81, 
114,  147 

epidermidis  (Lichen)  147 
erecta  (Parmelia)  232 
ericetorum  (Cetraria)  239 
erichsenii  (Verrucaria)  61 
erratica  (Lecidea)  21.  22,  29,  43.  55, 
81,  100*,  725,  167,  168,  177,  273 
erratica  (Lecidea  erratica  var.)  168 
erythrocardia  (  Physcia  ciliata  f. )  260 
erytlirocardia  ( Physcia  ciliata  var. ) 
260 

erythrocardia  (Physcia  obscura  var.) 
260 

Eupertusaria  (Pertusaria  subgenus) 
286 

evansii  (Cladonia)  82.  128,  156.  198. 
201,  243 

Evernia  241 

evolutoides  ( Stereocaulon  paschale 
var.)  178 

Fagus  13,  26.  28,  33,  47,  114.  119. 
155.  211,  215 

fallaciosa  (Polyblastiopsis)  148 
fallax  ( Arthopyrenia)  148 
fallax  (Physcia)  251 
fallax  (Polyblastiopsis)  148 
fallax  (Xanthoria)  30,  54,  55,  80, 
1 43,  218,  251,  275 

farinacea  (Cladonia)  128,  133,  190. 
195,  275 

farinacea  (Cladonia  furcata  y  scab- 
riuscula  f. )  195 


farinacea  (Ramalina)  141 
fastigiata  (Lichen  calcaris  var.)  243 
fastigiata  (Ramalina)  32,  58,  82,  142, 
243.  275 

fendleri  (Cetraria)  21,  82.  141.  238, 
239 

fendleri  (Platsyma)  238 
feracissima  (Caloplaca)  21,  55,  143, 

250 

fimbriata  (Cladonia)  80.  135,  188, 
189 

fimbriatus  (Lichen)  188 
flaccidum  (Collema)  160 
flavicans  (Lichen)  253 
flavicans  (Teloschistes)  83,  144,  253, 
276.  277 

flavovirescens  (Caloplaca)  21,  30,  55, 
81,  143,  250.  258 
flavovirescens  (Lichen)  250 
flexuosa  (Lecidea)  165 
floerkeana  (Cenomyce)  179 
floerkeana  (Cladonia)  81,  133,  179 
florida  (Usnea)  245 
floridana  (Cladonia)  82.  97*,  131, 
194,  271 

Foliosae  (Clausae  subsect.)  183 
frondifera  ( Parmelia  )  227 
furcata  (Cladonia)  55.  56,  60,  80, 
110,  128,  131.  196.  271,  273,  275 
furcatus  (Lichen)  196 
furfuracea  ( Pseudevernia )  73*,  82, 
87.  88.  122,  237.  281 
furfuraceus  (Lichen)  237 
fusca  (Anaptychia)  262 
fuscata  (Acarospora)  55,  60,  61,  81, 
99*.  114.  176,  205,  271,  272 
fuscatus  (Lichen)  205 
fuscidula  (Lecanora)  219.  220 
fuscorubella  (Bacidia)  172 

galbina  (Parmelia)  67,  81,  98*.  141, 
229,  231,  267,  271,  273,  275 
glabra  (Alectoria)  81.  141,  241,  244 
glabra  (Carya)  150 
glabrata  (Lecanora)  215 
glauca  (Smilax)  13 
glauconigrans  (Biatora)  171 
glauconigrans  (Catillaria)  81,  116, 

171 


INDEX  TO  NAMES 


323 


glaucum  (Leucobryum)  44 
grande  ( Rhizocarpon )  80,  117,  127, 

176 

grandidentata  (Populus)  25 
grandifolia  (Fagus)  13,  21,  25,  27, 
31,  49,  54,  59,  156,  210 
grandis  (Lecidea  petraea  var.  fuscoa- 
tra  f. )  176 

granulosa  (Lecidea)  42,  56.  81,  116, 
125,  165,  168,  169 
granulosa  (Lichen)  168 
Graphidaceae  158 
Grapliis  158,  288 
grayi  (Cladonia)  43,  188 
gregaria  (Arthonia)  151 
Gyalectaeeae  159 

Haematomma  82,  138.  223 
Haematomma  sp.  118,  138,  224,  273, 
275 

hageni  (Lecanora)  81,  138,  218 
liageni  (Lichen)  218 
halei  (Usnea  subfusca  var.)  247 
halodytes  ( Arthopyrenia)  146 
harmandi  ( Ochrolechia )  223 
Helopodium  ( Podostelides  series)  184 
herrei  (Cetraria)  240 
heterocliroa  (Anaptychia)  262 
hibernica  (Porina)  124,  156.  157, 
281,  289* 

hispida  (Physcia)  259 
horizontalis  (Peltigera)  164 
hossei  (Usnea)  246 
Hudsonia  106 

hueana  (  Physcia  orbicularis  f. )  260 

Hypogyninia  236 

Hypotrachyna  (Parmelia  subgenus) 
230,  23  1,  287 

hypotropa  (Parmelia)  56,  58,  59.  82, 
95*,  140.  176,  229,  230,  232,  233, 
273,  275 

hypotropoides  (Parmelia)  232 

Ilex  21.  31.  114,  119,  156 
ilicifolia  (Quercus)  10*,  12 
impexa  (Cladonia)  60,  87,  201 
impolita  (Arthonia)  150 
incana  (Byssus)  263 
incana  (Lepraria)  22,  118,  146,  263 


incrassata  (Cladonia)  49,  56,  57,  82, 
130,  131,  181,  273 
Insensibiles  (Eupertusaria  sect.)  210 
insignis  (Lecanora)  215,  217 
intermedia  ( Amelanchier)  13,  14,  59 
intermedia  (Bacidia)  127,  172,  174, 
175,  281 

intermedia  (Biatora  anomala  var.) 
173 

intermedium  (Rhizocarpon)  117,  127, 
111,  256 

inundata  (Bacidia)  81,  126,  175 
inundata  (Biatora)  175 
Fonaspis  213 
isidiata  (Parmelia)  228 
islandica  (Cetraria)  56,  60,  80.  202, 
239,  272  (See  also  "crispa”) 
islandicus  (Lichen)  239 

japonica  (Cladonia)  196 
jubata  (Alectoria)  241 
juniperina  (Cetraria)  240 
Juniperus  217,  226,  243.  253 

laciniata  (Parmelia  epiclada  var.)  231 
laciniata  (Parmelia  michauxiana  var.) 
231 

lactea  ( Polyblastiopsis)  148 
lacunosa  (Cetraria)  240 
laevigatum  (Nephroma)  76*.  82,  92*, 
120,  162,  269 

laevis  (Lecanora)  61,  137,  218.  219 
Lasallia  204 

lathamii  (Lecanora  caesiorubella 
subsp.)  32.  39*.  50,  54,  57,  82, 
111,  1 37,  214,  271 
lavata  (Ionaspis)  213 
Lecanora  115,  136,  213.  288,  289* 
Lecanora  (Lecanora  section  )  136 
Lecanoraceae  213 
Lecanora  sp.  137.  220 
Lecidea  115,  125,  165,  288.  289* 
Lecideaeeae  165 
leioplaca  (Pertusaria)  211 
Lepraria  146.  171,  263 
Lepraria  sp.  118,  146,  265 
leprarioides  (Haematomma)  224 
Leptcgium  119,  124,  160 
Leptorhaphis  147 


324 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


livida  (Parmelia)  58,  67,  81,  141, 

230.  231,  273 
Lobaria  40.  120,  124,  161 
Loculoascomycetidae  284,  287 
longissima  (Usnea)  81,  142,  244 
lurida  (Pannaria)  83,  119,  161,  275 
I uritl nm  (Collema)  161 

lusitana  (Parmelia)  228 
lutea  (Dimerella)  81,  116.  159 
luteola  (Bacidia)  172 
luteus  (Lichen)  159 
Lycopodium  16 

macilenta  (Cladonia)  56,  60,  81,  130, 
132.  180.  273 

macounii  (Melanaria)  115,  136,  212 
macrocarpa  (Lecidea)  80,  125,  169 
macrocarpa  (Patellaria)  169 
macrocarpon  (Vaccinium)  14,  16 
mammulata  (Gyrophora)  203 
mammulata  ( Lmbilicaria )  82,  135, 
203.  204 

marginata  ( Pertursaria )  207 
maritima  (Prunus)  10,  12,  13,  49,  58, 
238 

mateocyatha  (Cladonia)  82.  129,  134, 

186 

mediella  (Arthonia)  123,  150.  281 
Megaphyllae  (Chasmariae  series)  192 
melaena  (Lecidea)  152 
melaena  (Micarea)  81,  123,  126,  152 
Melanaria  212 
Melanoparmelia  235 
Melanotheca  155 
membranacea  (Crocynia)  265 
membranacea  (Lepraria)  265 
meroclilorophaea  (Cladonia)  188 
merrillii  (Usnea)  247 
mesomorpha  (Evernia)  81,  122,  241 
Micarea  115,  123,  152,  173,  176.  286 
michauxiana  (Parmelia)  82,  140,  141, 

231,  273 

michauxiana  (Parmelia  michauxiana 
var.)  231 
Micropliiale  159 

Microphyllae  (Chasmariae  series)  192 
microspora  (Verrucaria)  14*,  22.  29, 
58,  61,  83,  123,  153,  154,  155 
nricytho  (Lecidea)  168 


millegrana  (Physcia)  35,  37,  38*, 
39*,  40,  50.  54,  59,  81,  92,  103*, 
145,  225,  259.  261,  267,  274,  275 
milliaria  (Rinodina)  58,  81,  104*, 
144,  256,  257.  269.  274 
miniatum  ( Dermatocarpon)  80,  120, 
155 

miniatus  (Lichen)  155 
mitis  (Cladonia)  80,  1 29.  202,  203 
mitrula  (Cladonia)  184 
montagnei  (Ramalina)  243 
muhlenbergii  (Gyrophora)  204 
muhlenbergii  ( Umbilicaria)  82,  135, 
204.  276.  277 

multiformis  (Cladonia)  81,  134,  194 
multipuncta  (Pertusaria)  136,  206, 
207 

multipuncta  (Variolaria)  206 
muralis  (Lecanora)  81,  136,  219.  277 
muralis  (Lichen)  219 
muralis  (Verrucaria)  59,  80,  124,  154 
mutabilis  (Usnea)  82,  142,  245 
myriocarpoides  (Lecidea)  56,  125, 
169 

nearingii  (Lecidea)  167.  168 
nemoxyna  (Cladonia)  81.  135,  190, 
297 

nemoxynus  ( Baeomyces  radiatus  [3 
B.)  190 
Nephroma  162 
Nephroinaceae  111,  162 
nidulifera  (Alectoria)  21,  82,  95*, 
141,  242,  271,  272,  275 
niger  (Lichen)  161 

nigrescens  (Verrucaria)  59.  81,  124, 

154 

nigrum  ( Placynthium )  55,  80,  113, 
116,  161 

nitida  (Pyrenula)  54.  114,  155,  211, 
271,  274 

nitida  (Sphaeria)  155 
Nostoc  291* 

nucula  (Porina)  83,  118,  124,  157 

nylanderi  (Biatora)  169 

nylanderi  (Lecidea)  81.  126,  169,  170 

obscurata  (Anaptychia)  83,  145,  146, 

262 

obscurata  (Lecidea  petraea  y  L.)  177 


INDEX  TO  NAMES 


325 


obscurata  (Physcia)  262 
obscuratum  ( Rhizocarpon)  21,  22,  55, 
127,  167,  177 

ochrocarpia  (Cladonia  cristatella  f.) 
183 

ochrochlora  (Cladonia)  189 
Ochrolechia  115,  138,  211,  221,  299 
Ochrolechia  sp.  138,  223 
Ochroleucae  (Clausae  subsect.)  182 
ochroplwea  (Biatora)  223 
ochrophaeum  (Haematomma)  82, 
115,  119,  138,  223 
odora  (Gyalecta)  213 
odora  (Ionaspis)  115,  213,  281 
oligotropha  (Lecidea)  170 
olivaceum  (Coniosporium )  56 
olivetorum  (Parmelia)  72*,  82.  140, 
231,  232 

opaca  (Ilex)  25,  49,  57,  59,  91,  156, 
159 

Opegrapha  113,  123,  153,  287 
Opegraphaceae  153 
Opegrapha  sp.  118,  123 
opegraphella  (Xylographa)  82.  113, 

158 

orbicularis  (Lichen)  260 
orbicularis  (Physcia)  35.  50,  59,  81, 
121.  145,  260,  261,  274 
orbicularis  (Physcia  orbicularis  f.) 
260.  261 

oregonensis  (Ochrolechia)  222 
oreina  (Lecanora  straminea  (1  L.) 
257 

oreina  (Rinodina)  21,  55.  80.  116, 
144,  225,  257 

orphninum  (Rhizocarpon)  177 
Osmunda  16 

oxycoccos  (Vaccinium)  14,  16 

pachysperma  (Rinodina)  105*,  145, 
257.  269,  274 

pallescens  (Ochrolechia)  221.  222, 
223 

pallida  (Lecanora)  88,  214 
palmatula  (Anaptychia)  263 
palmulata  (Anaptychia)  82.  145,  146, 
262,  275 

palmulata  (Psoroma)  262 
palustris  (Pinus)  12 

Pannaria  161 


Pannariaceae  161 

papillaria  (Lichen)  178 
papillaria  (Pycnothelia)  28,  42.  56, 
82,  103*,  116,  122,  127,  178,  179, 
268,  274 

papulosa  (Gyrophora)  204 
papulosa  (Umbilicaria)  82,  87.  88, 
135,  204 

parasema  (Buellia)  255 
parasitica  (Cladonia)  49,  56,  57,  81, 
130,  132,  192,  193.  273 
parasitica  (Lecanora  scruposa  var. ) 
159 

parasiticus  ( Diploschistes  scruposus 
var. )  159 

parasiticus  (Lichen)  192 
parella  (Ochrolechia)  57,  96*,  138, 
221.  281 

parellus  (Lichen)  221 
parietina  (Xanthoria)  30,  32,  54,  59, 
82,  92,  105*,  110,  143,  218,  251, 
252,  269,  275 
parietinus  (Lichen)  251 
Parmelia  40,  121,  139,  166.  227,  240 
Parmeliaceae  226 
Parnieliopsis  121,  139,  226 
paschale  (Stereocaulon )  178 
Patellaria  169 

Peltigera  119,  124,  163,  294 
Pcltigeraceae  163 

pensylvanica  (Myrica)  10*,  12,  13, 
14,  49.  58.  146 

perforata  (Parmelia)  54,  58.  81,  98*, 
140,  229,  230,  232,  267,  271,  274, 
275 

perforalus  (Lichen)  232 

perlata  (Parmelia)  83.  140,  229.  230. 

233 

perlatus  (Lichen)  233 
pertusa  (Pertusaria)  210.  212 
Pertusaria  53,  114,  115,  135,  206,  285, 
286,  287,  290,  292 
Pertusariaceae  206 
Perviae  (Cladonia  sect.)  192 
phaeocephala  ( Chaenotheca)  22,  81, 
113,  117,  153 

phaeoceplialus  (Lichen)  153 

Phaeograpliis  158 

Physcia  53,  65,  1 21,  145,  172.  258 
Pliysciaceae  111,  253 


326 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


physodes  (Hypogymnia)  35,  54.  56, 
80,  120,  237,  240,  273,  288 
physodes  (Lichen)  237 
Picea  33 

piedmontensis  (Cladonia)  82,  132, 

183,  191 

pinastri  (Lecanora)  215 

pinicola  ( Arthopyrenia )  122,  146 

pinicolci  (Pyrenula  punctiformis  var. 

cineropruinose  f. )  146 
Pinus  26,  33 

pityrea  (Capitularia)  191 
pityrea  (Cladonia)  83,  133,  191 
Placodium  (Lecanora  sect.)  136 
placorodia  (Parmelia)  226 
placorodia  (  Parmeliopsis )  21,  45,  49, 
82.  96*.  139.  226,  227,  238,  267, 
270,  271.  274 
Placynthium  161 
planetica  (Lecidea)  168 
planetica  (Lecidea  erratica  var.)  168 
platycarpa  (Lecidea)  169 
Platysma  (Cetraria  subgenus)  292 
Pleurococcus  40 
pleurota  (Capitularia)  182 
pleurota  (Cladonia)  55,  56,  80,  99*, 
134,  182,  183,  268,  273 
plicatile  (Rhizocarpon )  127,  177,  178 
plicatilis  (Lecidea)  177 
plittii  (Parmelia)  83,  1 39,  228.  233 
Podostelides  (Clausae  subsect.)  184 
Polyblastiopsis  147 
polycarpa  (Cladonia)  184 
polycarpa  (Xanthoria)  252 
polydactyla  (Peltigera)  80.  124,  164, 
275 

polydactyla  (Peltigera  polydactyla 
var.)  164 

polydactylon  (Lichen)  164 
polymorpha  (Arthonia)  123,  150 
polyspora  (Buellia)  81,  144,  254,  272 
polyspora  (Buellia  myriocarpa  var.) 
254 

Polytrichum  44 
ponderosa  (Pinus)  170,  227 
populifolia  (Betula)  21,  57,  147 
Porina  114,  124,  156 
Porinaceae  156 

praetextata  (Peltigera)  80.  124,  164. 
165,  276 


praetextata  ( Peltidea  ulorrhiza  var.) 
164 

prasina  (Biatora)  152 

prasina  (Catillaria)  152 

prasina  (Micarea)  22,  56,  81.  123, 

152 

prinus  (Quercus)  13,  25,  49.  50.  51 
privigna  ( Lecidea )  205 
privigna  (Sarcogyne)  135,  205 
propinqua  (Pertusaria)  82,  135,  207 
pruinosa  (Arthonia)  150 
pruinosa  (Sarcogyne)  205 
prunastri  (Evernia)  241 
Pseudevernia  237.  288 
pseudoacacia  (Robinia)  25.  55 
pseudospeciosa  (Anaptychia)  83.  145, 
146,  262,  263 

pulmonaria  (Lobaria)  34,  57,  81,  92*, 
124,  161,  162,  237,  269,  275,  276, 
277 

pulmonarius  (Lichen)  161 
punctata  (Buellia)  81,  144,  254 
punctata  (Verrucaria)  254 
punctiformis  (Arthonia)  123,  150, 
151 

purpurea  (Sarracenia)  16 
pusilla  (Schizaea)  201 
pustulata  (Pertusaria)  206,  212 
pustulata  (Umbilicaria)  87,  88 
Pycnothelia  178,  292 
pyracea  (Caloplaca)  30.  80.  143,  248, 
249.  250,  251.  276,  277 
pyracea  (Parmelia  cerina  var.)  250 
Pyrenula  155 
Pyrenulaceae  155 

pyxidata  (Cladonia)  43,  60,  80.  129, 
134,  186,  187,  275 
pyxidatus  (Lichen)  186 

Pyxine  258 

quercicola  ( Polybastiopsis)  114,  147, 
148*.  149*,  281.  289* 
quercina  (Parmelia)  229 
quercizans  (Lobaria)  34.  57.  67.  82, 
87.  88,  92*.  124,  162,  237,  269,  275 
Quercus  33,  153,  155 
quinquefolia  ( Parthenosissus)  13 

racemosa  (Cladonia  furcata  f. )  56 
radiata  (Arthonia)  151 


INDEX  TO  NAMES 


327 


radicans  (Toxicodendron)  12 
Ramalina  107,  122,  141,  242,  288 
rangiferina  (Cladonia)  80.  129,  201, 
276,  277 

rangiferina  (Cladonia  rangiferina 
subsp.  rangiferina  var.)  201 
rangiferinus  (Lichen)  201 
reductum  ( Rhizocarpon  obscuratum 
f.)  Ill,  177 

reticulata  (Parmelia)  81.  140,  230, 
233,  275 

rhaphidosperma  (Porina)  156,  157 
Rhizocarpon  115,  127,  176 
rigida  (Pinus)  10*,  12,  13,  14*,  21, 
24,  27,  28,  30,  49,  56.  166,  170, 
203,  226,  227,  237,  238,  240,  242, 
267,  275 

Rinodina  115,  144,  256 
robbinsii  (Cladonia)  130,  132,  183 
Robinia  30 

robusta  (Parmelia)  229 
rosella  (Lecanora  pallescens  var.)  222 
rosella  (Ochrolechia)  138,  221,  222, 
223 

rosella  (Pertusaria  pallescens  var.) 
22 1 

roseus  (Baeomyces)  22,  28,  30.  42. 

56.  82.  98*,  116,  122,  179,  268.  272 
rubella  (Bacidia)  172 
rubescens  (Pertusaria)  207 
rubina  (Lecanora)  80,  120,  136,  219 
rubinus  (Lichen)  219 
rubra  x  coccinea  (Quercus)  19 
rubra  (Quercus)  5,  13,  19,  21,  24.  26. 

27,  28,  50,  51,  52.  53 
rubropulchra  (Physcia  orbicularis  f . ) 
260,  261 

rubrum  (Acer)  13,  14,  16.  25,  27,  49. 
54.  57.  59.  91.  162,  170.  175.  208. 
210,  223,  233,  275 
Rubus  13 

rudecta  (Parmelia)  21,  35,  50.  54,  57. 
81,  139,  234,  274 

rufescens  (Lichen  caninus  var.)  163 
rufescens  (Opegrapha)  119,  123,  153 
rufescens  (Peltigera)  110 
rufescens  (Peltigera  canina  var.)  125, 
163,  164,  165 

rupestris  (Synechoblastus)  160 
rupestris  (Verrucaria)  154 


saccata  (Solorina)  80,  120,  163 
saccatus  (Lichen)  163 
saccharinum  (Acer)  170 
saligna  (Lecanora)  174 
salina  (Rinodina)  59,  144.  258 
santensis  (Cladonia)  57,  82,  94*,  132, 
134,  156,  193 

Sarcogyne  114,  135,  204,  225 
sarcopsis  (Lecanora)  220 
saxatile  (Stereocaulon )  81,  122,  178. 
179,  281 

saxatile  (Stereocaulon  evolutoides 
var.)  179 

saxatilis  (Lichen)  234 

saxatilis  (Parmelia)  35,  50,  54,  77, 

80,  140,  234,  236,  274 
saxicola  (Bacidia  trisepta  f . )  175 
scabriuscula  (Cenomyce)  195 
scabriuscula  (Cladonia)  80,  128,  133, 

195,  275 

scalaris  (Lecidea)  21,  45,  49,  81,  96*, 
117,  125,  130,  170.  267,  271.  273 
scalaris  (Lichen)  170 
scalaris  (Psora)  170 
schweinitzii  (Bacidia)  81,  126,  175 
schweinitzii  (Biatora)  175 
scoparius  (Andropogon)  13,  14 
scripta  (Graphis),  21,  35.  50,  54,  57, 

81.  100*,  113,  158,  267,  271,  273, 
275.  291 

scriptus  (Lichen)  158 

scruposus  ( Diploschistes)  80,  114, 

159 

scruposus  ( Diploschistes  scruposus 
var.)  159 

scruposus  (Lichen)  159 
serotina  (Prunus)  10*,  13,  14,  25.  49, 
59 

sexlocularis  (Arthonia)  123,  151.  281 
siderea  (Arthonia)  102*,  123,  150, 
151,  272 

silicicola  (Verrucaria)  22.  29.  61,  83, 
105*,  124,  154.  276 
simplex  (Lichen)  205 
simplex  (Sarcogyne)  80.  135.  205 
simulata  (Cladonia)  82.  131,  191 
Smilax  13 
Solorina  163 

sordidescens  (Lecidea)  152 


328 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


sordidescens  (Micarea  prasina  var.) 

152 

sorediata  (Lecidea)  258 
sorediata  (Pyxine)  82,  121,  145,  258 
sorediifera  (Anaptychia)  262 
speciosa  (Anaptychia)  262,  263 
Sphagnum,  14,  16 

spuria  (Peltigera  canina  var.)  124, 

164 

spurius  (Lichen)  164 
squamosa  (Cladonia)  60,  80,  98*, 
109,  130,  134 ,  193,  194,  268 
squamosus  (Lichen)  193 
squamulifera  (Cladonia  pityrea  var. 

zwackii  f. )  191 
stellaris  (Lichen)  261 
stellaris  (Physcia)  54,  69*,  81,  97*, 
145,  261,  267,  270.  271,  274 
stellata  (Quercus)  148 
stenophylla  (Parmelia)  55,  81,  139, 
234.  235,  236,  275 

stenophylla  (Parmelia  conspersa  f} 
P.)  234 

stenospora  (Ramalina)  82,  142,  243, 
276,  277 

Stereocaulaceae  178 
Stereocanlon  178,  292 

steriza  (Lecidea)  169 

steriza  (Lecidea  confluens  y  L.)  169 

Sticta  162 
Stictaceae  161 

stigmaea  (Buellia)  55,  82,  117,  144 , 
176,  255,  256 

stillingiana  (Buellia)  59,  81,  118,  144, 
253,  255,  271.  272 

Stramineoflavidae  (Cocciferae  series) 

181 

strepsilis  (Baeomyces)  183 
strepsilis  (Cladonia)  42,  56,  60,  81, 
103*,  130,  132,  183,  268,  271,  273, 
299 

striata  (Ochrolechia  parella  f. )  221 
strigosa  (Usnea)  32,  36,  50,  58,  59, 
82.  142.  245,  246,  247,  274,  275 
strigosa  (Usnea  florida  y  U.)  245 
subacuta  (Cladonia  pityrea  var. 
zwackii  f. )  191 

subaurifera  (Parmelia)  21,  50.  54,  59, 
82,  121,  139,  235.  271,  274 


subcariosa  (Cladonia)  42,  56,  81, 
109.  130,  132,  184,  185 
subfurvum  (Collema)  67,  82,  92*, 
119,  160,  275,  276,  277 
subfurvus  ( Synechoblastus  flaccidus 
var.)  160 

subfusca  group  (Lecanora)  215,  220 
subfusca  sensu  Motyka  (Usnea)  56, 
142,  246,  247 

subfusca  Stirt.  (Usnea)  247 
subfuscata  (Lecanora)  215,  220 
Subglaucescentes  (Cocciferae  series) 

179 

subintricata  (Lecanora)  45,  137,  219, 
220 

subintricata  (Lecanora  varia  var.)  219 
sublitoralis  (Arthopyrenia)  146 
submitis  (Cladonia)  10*.  22,  30.  32, 
56,  60,  82,  104*.  129,  199,  202,  203, 
239,  268,  271.  273,  275,  298 
subpertusa  (Pertusaria)  136,  207, 

208*,  209*,  210,  281 
subrudecta  (Parmelia)  82,  101*,  139, 
235, 274 

subrugosa  (Lecanora)  215 
subrangiformis  (Cladonia)  196 
substraminea  (Cladonia)  183 
subsuperficialis  ( Verrucaria)  153,  154 
subtenuis  (Cladonia)  44,  55,  81,  87, 
129,  166,  199,  200,  202,  271,  273, 
275 

subtenuis  (Cladonia  subtenuis  f.)  199, 
200 

subtenuis  (Cladonia  tenuis*  Cl.)  199 
subtilis  (Physcia)  82,  145,  260.  261, 
262 

subtrichodea  (Usnea  hossei  f. )  246 
sulcata  (Parmelia)  21,  32,  35.  37, 
38*,  39*.  40,  50,  54,  58.  59.  77.  80, 
109,  140.  234,  236,  274,  275 
sylvatica  (Cladonia)  42,  60.  202,  276 
sylvatica  (Nyssa)  14,  16 
sylvicola  (Lecidea)  168 
symmicta  (Lecanora)  80.  126,  138, 
217.  220 

symmicta  (Lecanora  varia  7  L.)  220 

taeda  (Pinus)  12,  45 
tacdiosa  (Arthonia)  151 


INDEX  TO  NAMES 


329 


taediosum  (Arthothelium)  114,  151, 
152 

tartarea  (Ochrolechia)  222 
tasmanica  (Parmelia)  139,  235,  236 
Teloschistaceae  111,  248.  292,  298 
Teloschistes  122,  144,  252 
Tenues  (Cladonia  sect.)  199 
tenuis  (Cladonia)  60,  87 
terrae-novae  (Cladonia)  82,  83,  87, 
92*,  129,  198,  200,  201 
tetrathalamia  (Pertusaria)  211 
Thallostelides  (Clausae  subsect.)  185 
thamnodes  (Evernia  prunastri  var. ) 

241 

Thuja  172 

thyoides  (Chamaecyparis)  14,  25,  49, 
50,  56,  153,  173,  226,  237,  238, 
242,  247,  267,  276 
tomentosa  (Carya)  9,  106,  207 
tomentosa  (Hudsonia)  10*,  12,  28. 

242 

torquata  (Pertusaria)  207 
trachythallina  (Pertusaria)  54,  118. 

136.  210,  224,  270,  271,  274 
Trebouxia  115,  120,  126,  150,  175, 
291*,  294 

tremelloides  (Leptogium)  160 
tremuloides  (Populus)  172 
Trentepohlia  1  15,  118,  122,  126,  148. 

150,  157,  175,  291* 
tribacia  (Physcia)  259 
tribacoides  (Physcia)  82,  145,  262 
trichodea  (Usnea)  56,  57,  82,  95*, 
142.  246,  247,  267,  271,  274 
trisepta  (Bacidia)  123,  126,  175 
trisepta  (Lecidea)  175 
Trypethelium  156 

tuberculifera  (Pertusaria)  83,  136, 

210,  211,  270,  271,  274,  275,  281 
tuckermanii  (Cetraria)  82,  121,  141. 
239.  240,  275 

tulipifera  (Liriodendron)  5,  9 
turgescens  (Buellia)  81,  101*,  144, 

255 

typicct  (Peltigera  aphthosa  var.)  163 
typica  (Peltigera  polydactyla  var.) 
164 


uliginosa  (Biatora)  60 
uliginosa  (Lecidea)  28,  42,  44,  56,  60, 
81,  116,  125,  170,  271,  273 
uliginosus  (Lichen)  170 
Ulmus  26,  30,  218 
ulorrhiza  (Peltidea)  164 
ulorrhiza  (Peltigera  canina  var.)  125, 
164 

Umbilicaria  120,  135,  203,  287,  294 
Umbilicariaceae  203 
umbrina  (Bacidia)  61,  81,  126,  176 
umbrina  (Lecidea)  176 
Unciales  ( Perviae  subsect.)  128,  197 
uncialis  (Cladonia)  44,  56,  60.  80, 
104*,  128,  190,  197,  198,  268.  271, 
273,  275 

uncialis  (Lichen)  197 
Urceolaria  159 

Usnea  122,  142,  244,  284,  285 
Usneaceae  241 
Usnea  sp.  142,  246 
Utricularia  16 

uva-ursi  ( Arctostaphylos)  10*,  12,  28. 
42 

vaccilans  (Vaccinium)  12 
Vaccinium  13,  14* 
varia  (Lecanora)  81.  138,  220 
varians  (Lecidea)  58,  59,  97*,  126. 
171.  271,  273 

variolosa  (Peltigera  aphthosa  f. )  163 
variolosa  (Peltigera  aphthosa  var.) 
163 

varius  (Lichen)  220 
velata  (Parmelia)  211 
velata  (Pertusaria)  82,  92*,  136,  211 
velutina  (Quercus)  5,  9,  13,  14*,  19, 
21,  24,  26,  49,  50,  51,  52,  53,  54, 
210,  229,  233,  234,  236.  241,  254, 
262 

vernalis  (Lecidea)  80,  126,  171.  220 
vernalis  (Lichen)  171 
vernix  (Toxicodendron)  14 
Verrucaria  114,  123,  153 
Verrucariaceae  153 
verticillata  (Cladonia)  80.  134,  185, 
186,  275,  294 

verticillata  (Cladonia  pyxidata*  C.) 
185 

verticillata  (Ilex)  25.  49.  57,  59,  107 


330 


THE  UNIVERSITY  OF  THE  STATE  OF  NEW  YORK 


virella  (Physcia)  260 
virens  (Trypethelium  )  21,  31,  54,  57, 
59,  82,  114,  119,  156 
virginiana  (Juniperus)  25.  49.  59, 
241.  243,  244,  260,  261 
virginica  (Woodwardia)  14 
viridescens  (Lecidea)  81,  118,  126, 
171 

viridescens  (Lichen)  171 
viridis  (Cetraria)  56.  57,  82,  93*, 
120,  141,  240.  267.  271,  272 
viridis  ( Protococcus )  22 
vitellina  (Candelariella)  21,  59.  80. 

102*,  116,  138.  225 
vitellina  (Candelariella  vitellina  var. ) 
225 

vitellinus  (Lichen)  225 


vulcanica  (Cladonia)  57,  83,  109, 
130,  132,  180.  181 
vulgaris  (Calluna)  42 
vulgata  (Opegrapha)  153 

willeyi  (Ramalina)  59.  74*,  82,  141, 
243,  244.  276 

xanthodes  (Pertusaria)  32,  35,  50,  57, 
59,  82.  136,  211,  212.  274 
Xanthoria  25,  30.  120,  143,  172,  251 
Xanthoparmelia  228,  229 
Xylographa  158 

z onata  (Crocynia)  263 
zonata  (Lepraria)  55.  117,  146,  263, 
264*,  281 

zwackii  (Cladonia  pityrea  var.)  191 


Text  set  in  Linotype  Times  Roman 
Heads  set  in  Garamond 


UK587  ,B7  c  2  -  v 

Br°MC(f|l|||||inMi/Im1  Mchens  of  Lon9  ilia" 


3  5185  5o,i"2I0,I"I59I37 


M  724-Mr  66-2200